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Effects of water stress cycles on turgor maintenance processes in pear leaves (Pyrus communis).

Effects of water deficits on leaf turgor maintenance processes were analyzed for pear trees (Pyrus communis L. cv. "Barlett") grown in 120-liter containers. Four irrigation treatments were applied: a well-watered control treatment, a spring water stress cycle (Sp), a summer water stress cycle (Su), and a spring plus summer water stress cycle (Sp + Su). For the Sp treatment, water application was progressively reduced from 100 to 20% of the control dose over a period of 27 days in spring. For the Su treatment, water application was progressively reduced over 23 days in summer, from 100 to 20% of the control dose. The Sp + Su treatment comprised both the spring and summer drought stress cycles. Pressure-volume (P-V) curves were constructed and stomatal conductances were determined for pear leaves from each treatment during the spring and summer stress cycles. Leaf water potential (Psi(pi) (0)) and relative water content (R(0)) at the turgor loss point of control leaves tended to decrease from spring to summer. Changes in leaf osmotic water potential at full turgor (Psi(pi) (100)) and in symplast water fraction (R(s)) did not explain the seasonal decrease in Psi(pi) (0). The water stress treatments had no effect on Psi(pi) (100), but R(s) was reduced by the water stress treatments, particularly during the summer stress cycle of the Su and Sp + Su treatments. The decrease in R(s) was correlated with an increase in the slope of the linear region of the P-V curve. Such a coupled adjustment would lead to increased water uptake capacity of water-stressed trees only under non-turgor conditions. Furthermore, pear leaves did not actively accumulate solutes. We conclude, therefore, that changes in leaf tissue water relations as a result of leaf acclimation to water stress are unlikely to facilitate maintenance of fruit productivity under drought.

Journal Article↗

Effects of cone-induction treatments on black spruce (Picea mariana) current-year needle development and gas exchange properties.

Both drought and root pruning (RP) increased the number of cones induced when black spruce (Picea mariana (Mill.) B.S.P.) grafts were injected with gibberellins A(4/7) (GA), but their effects on predawn shoot water potential and current-year needle development differed. Drought decreased predawn shoot water potential (Psi(pd)), but only during the period when irrigation was withheld, and it had no effect on the growth or gas exchange properties of current-year needles. Conversely, root pruning had little effect on Psi(pd), but it resulted in trees with smaller current-year needles that had lower nitrogen and chlorophyll concentrations and reduced rates of gas exchange up to the later stages of shoot elongation compared with needles of control trees. These findings are discussed in relation to potential effects on the development of induced cones in the following growth cycle.

Journal Article↗

Stomatal conductance, growth and root signaling in Betula pendula seedlings subjected to partial soil drying.

Seedlings of Betula pendula Roth were grown with their root systems separated between two soil compartments. Four treatments were imposed: (i) adequate irrigation in both compartments (WW, controls); (ii) adequate irrigation in one compartment and drought in the other compartment (WD); (iii) drought in both compartments (DD); and (iv) half of the root system severed and the remainder kept well-watered (root excision, RE). Predawn leaf water potential, stomatal conductance, soil-to-leaf specific hydraulic conductance, and root and leaf growth decreased in DD-treated seedlings, which also displayed severe leaf shedding (30% loss in leaf area). The DD treatment also resulted in increased concentrations of abscisic acid (ABA) and its glucose ester in the xylem sap of roots and shoots compared to concentrations in control seedlings (about 200 versus 20 nM). Despite the difference in xylem sap concentrations, total ABA flux to the shoots was similar in the two treatments (1-2 pmol ABA m(-2) leaf area s(-1)) as a result of reduced transpiration in the DD-treated seedlings. Compared with root growth in control plants, root growth increased in the RE-treated plants and decreased in the drying compartment of the WD treatment; however, the RE and WD treatments only slightly reduced leaf expansion, and had no detectable effects on shoot water relations or ABA concentrations of the root and shoot xylem sap. We conclude that short-term soil water depletion affecting only 50% of the root system does not cause a measurable stress response in birch shoots, despite root growth cessation in the fraction of drying soil.

Journal Article↗

A mathematical and statistical analysis of the curves illustrating vulnerability of xylem to cavitation.

When vulnerability curves are used to assess the susceptibility of plants to drought, the water potential corresponding to 50% loss of conductivity is frequently used as a measure of susceptibility. However, this value does not distinguish between different patterns of conductivity loss, such as a rapid loss over a narrow water potential range versus a gradual decline in conductivity. We have applied an exponential sigmoidal equation to vulnerability curves obtained from four closely related Eucalyptus clones differing in drought tolerance. The coefficients of the equation were evaluated and statictically compared among the clones. If the air-seedling hypothesis of cavitation is accepted, these coefficients have biological relevance. One of the coefficients describes the position of the curve on the water potential axis and is equivalent to the water potential corresponding to 50% loss of conductivity. This coefficient could reflect the size of the largest pit pore per xylem vessel. Another coefficient is related to the slope of the conductivity loss, and could reflect the range in maximum pit pore size per vessel.

Journal Article↗

Diurnal changes in water conduction in loblolly pine (Pinus taeda) and Virginia pine (P. virginiana) during soil dehydration.

We studied diurnal changes in water conduction during soil dehydration in 37-month-old seedlings of one Virginia pine (Pinus virginiana Mill.) and two loblolly pine (P. taeda L.) sources, one from North Carolina (NC) and the other from the "Lost Pines" areas of Texas (TX), in an environmentally controlled growth chamber. For seedlings of similar biomass, the TX source had higher values of transpiration, needle conductance, and plant hydraulic conductivity under well-watered conditions than the NC source. Under dry soil conditions, the TX source had lower values of water conduction than the NC source. The Virginia pine source responded similarly to the TX source under both well-watered and dry soil conditions. For all three pine sources, gradients between soil and needle water potentials were greatest when the seedlings were moderately stressed. The TX and Virginia pine sources had higher gradients and lower daytime needle water potentials under moderate stress conditions than the NC source. Predawn needle water potentials did not differ among the pine sources. We conclude that the TX and Virginia pine sources have decreased daytime needle water potentials and increased water potential gradients during the daytime under moderate stress conditions, but with no disruption of recovery at predawn. The greater rates of transpiration and water conduction by the TX source compared with the NC source under well-watered conditions suggest a means by which growth can be maximized prior to the onset of drought, thereby enhancing survival of loblolly pines in drought-prone environments.

Journal Article↗

Influence of weather on cork-ring width.

Ring-width series of cork from Quercus suber L. trees growing at two sites in Extremadura (southwestern Spain) were analyzed in relation to monthly precipitation and temperature, and to climatic indices combining both variables. Ring width of cork showed strong positive correlations with precipitation, especially during the fall and winter. Moderately low temperatures were favorable for cork growth, except in winter and during the onset of phellogen activity. We conclude that drought or temperature, or both, can limit cork growth during the annual drought period.

Rain↗

Quercus species differ in water and nutrient characteristics in a resource-limited fall-line sandhill habitat.

We compared co-occurring mature Quercus laevis Walt. (turkey oak), Q. margaretta Ashe (sand post oak) and Q. incana Bartr. (bluejack oak) trees growing in resource-limited sandhill habitats of the southeastern United States for water and nutrient characteristics. The Quercus spp. differed in their distribution along soil water and nutrient gradients, and in their access to and use of water, even though the study year was wetter than average with no mid-season drought. Quercus laevis had the greatest access to soil water (least negative pre-dawn water potential, psi(pd)) and the most conservative water-use strategy based on its relatively low stomatal conductance (g(s)), high instantaneous water-use efficiency (WUE), least negative midday water potential (psy(md)) and high leaf specific hydraulic conductance (K(L)). Quercus margaretta had the least conservative water-use characteristics, exhibiting relatively high g(s), low instantaneous WUE, most negative psi(md), and low K(L). Quercus margaretta also had a low photosynthetic nitrogen-use efficiency (PNUE), but a high leaf phosphorus concentration. Quercus incana had the poorest access to soil water, but intermediate water-use characteristics and leaf nutrient characteristics more similar to those of Q. laevis. There were no species differences for photosynthesis (A), leaf nitrogen on an area basis, or seasonally integrated WUE (delta13C). Both A and g(s) were positively correlated for each species, but A and g(s) were generally not correlated with psi(pd), psi(md) or delta psi(pd-md). Although we found differences in resource use and resource status among these sandhill Quercus spp., the results are consistent with the interpretation that they are generally drought avoiders. Quercus laevis may have an advantage on xeric ridges because of its greater ability to access soil water and use it more conservatively compared with the other Quercus spp.

Environment↗

Leaf water relations and stomatal behavior of four allopatric Eucalyptus species planted in Mediterranean southwestern Australia.

In 1986, four allopatric Eucalyptus species (E. camaldulensis Dehnh, E. saligna Smith, E. leucoxylon F. Muell and E. platypus Hook.) were planted together in a 480-mm rainfall zone, in 8-m wide contour belts as part of a plan to minimize waterlogging and secondary salinization. Throughout 1997, 1998 and 1999, there was significant inter-specific variation in predawn leaf water potential (Psi(pd)); however, maximum stomatal conductance (g(sm)) only differed significantly between species in mid to late summer. Relationships between g(sm) and Psi(pd) were significant and showed that stomata of E. camaldulensis were significantly more sensitive to Psi(pd), and presumably soil water potential, than stomata of E. leucoxylon or E. platypus. When applied to the Psi(pd) data, these relationships predicted that g(sm), and by inference transpiration, varied much less between species than Psi(pd). Diurnal measurements throughout the season confirmed this prediction, and showed that E. camaldulensis and E. saligna avoided drought by gaining access to deeper water, whereas E. leucoxylon and E. platypus maintained greater g(sm) at a given water stress than E. camaldulensis or E. saligna. Osmotic potentials measured after rehydration and water release curves of the leaves indicated that different mechanisms accounted for the apparent drought tolerance of E. leucoxylon and E. platypus. In summer, E. leucoxylon reduced osmotic potential at full and zero turgor by similar amounts compared with winter. In summer, E. platypus had a significantly lower bulk elastic modulus and relative water content at turgor loss point than E. camaldulensis, E. saligna or E. leucoxylon. This elastic adjustment resulted in a larger difference between osmotic potential at full and zero turgor in summer than in winter. The inherently low osmotic potential in E. leucoxylon and elastic adjustment in E. platypus resulted in turgor loss at a similar and significantly lower water potential than in E. camaldulensis or E. saligna. These results have implications for species selection for planting to manage groundwater recharge in areas prone to waterlogging and secondary salinization.

Journal Article↗

Effects of nutrition and soil water availability on water use in a Norway spruce stand.

We investigated effects of nutrition and soil water availability on sap flux density, transpiration per unit leaf area (EL), and canopy stomatal conductance (GS) of Norway spruce (Picea abies L. (Karst.)) in northern Sweden during the 1996 growing season. Our objectives were to determine (1) if artificially imposed drought (65% rain diversion) reduces soil water sufficiently to cause physiological limitations to whole-tree and plot-scale water transport, and (2) whether increased capacity for water transport resulting from fertilization-induced increases in leaf (> 3-fold) and sapwood areas (> 2.3-fold) deplete soil water sufficiently to cause a negative feedback on GS and EL. We monitored soil water content (theta) and soil water potential (PsiS) in control (C), drought (D), fertilized (F) and irrigated + fertilized (IL) treatment plots, along with site meteorological conditions. Ten trees per plot were monitored for sap flow. Although there were significant treatment differences in mean daily EL (C > D > F; P < 0.01) and GS (C > D > F; P < 0.05), variation in absolute magnitudes was small. Therefore, transpiration differences on a unit ground area basis (EC) were nearly proportional to leaf area differences. Precipitation was well distributed throughout the study period and so PsiS remained high, except during short dry periods in Plot F when it declined rapidly. Thus, although soil water was not limiting to GS, EL or EC when precipitation was uniformly distributed throughout the growing season, we cannot conclude that water availability would not limit GS in fertilized stands if the seasonal distribution of precipitation were altered.

Picea↗

Gas exchange characteristics of a Canarian laurel forest tree species (Laurus azorica) in relation to environmental conditions and leaf canopy position.

Diurnal courses of gas exchange were measured over a 1-year period in fully expanded current-year leaves in the upper (sun-exposed, 18 m above ground) and the lower (shaded, 12 m above ground) canopy of Laurus azorica (Seub.) Franco, a major canopy species of the Canarian laurel forest in Tenerife, Canary Islands, Spain. Laurus azorica exhibited high leaf plasticity in gas exchange characteristics, with a maximum carbon assimilation rate (Amax) of shade leaves about 50% that of sun leaves. This difference reflects the high leaf area index (LAI) of the stand and the correspondingly sharp light attenuation with increasing canopy depth. In sun leaves, Amax peaked at about 11 micromol m-2 s-1 and maximum transpiration (E) was about 8 mmol m-2 s-1, which corresponded with a maximum stomatal conductance (gs) of about 650 mmol m-2 s-1. Mean maximum instantaneous water-use efficiency (WUE) was 1.5 mmol mol-1 and the mean maximum A/gs was 20-35 micromol mol-1. Mean minimum internal CO2 concentration (Ci) was 225 micromol mol-1. Although high air vapor pressure deficit (VPD) caused a small decrease in gs, it remained high enough to maintain relatively high A and E. These gas exchange characteristics indicate a non-conservative use of water, which is appropriate for a species subject to droughts that are mild or of short duration. In this respect, Laurus azorica differs from its congener, L. nobilis L., of the Mediterranean region and other shrubs growing in Mediterranean-type climates in California and Chile that have to withstand more severe or more prolonged droughts.

Atlantic Islands↗

Carbon dioxide and water vapor exchange by young and old ponderosa pine ecosystems during a dry summer.

We investigated key factors controlling mass and energy exchange by a young (6-year-old) ponderosa pine (Pinus ponderosa Laws.) plantation on the west side of the Sierra Nevada Mountains and an old-growth ponderosa pine forest (mix of 45- and 250-year-old trees) on the east side of the Cascade Mountains, from June through September 1997. At both sites, we operated eddy covariance systems above the canopy to measure net ecosystem exchange of carbon dioxide and water vapor, and made concurrent meteorological and ecophysiological measurements. Our objective was to understand and compare the controls on ecosystem processes in these two forests. Precipitation is much higher in the young plantation than in the old-growth forest (1660 versus 550 mm year-1), although both forests experienced decreasing soil water availability and increasing vapor pressure deficits (D) as the summer of 1997 progressed. As a result, drought stress increased at both sites during this period, and changes in D strongly influenced ecosystem conductance and net carbon uptake. Ecosystem conductance for a given D was higher in the young pine plantation than in the old-growth forest, but decreased dramatically following several days of high D in late summer, possibly because of xylem cavitation. Net CO2 exchange generally decreased with conductance at both sites, although values were roughly twice as high at the young site. Simulations with the 3-PG model, which included the effect of tree age on fluxes, suggest that, during the fall through spring period, milder temperatures and ample water availability at the young site provide better conditions for photosynthesis than at the old pine site. Thus, over the long-term, the young site can carry more leaf area, and the climatic conditions between fall and spring offset the more severe limitations imposed by summer drought.

California↗

Physiological traits of two Populus x euramericana clones, Luisa Avanzo and Dorskamp, during a water stress and re-watering cycle.

We compared responses to drought and re-watering of greenhouse-grown cuttings of Populus x euramericana (Dode) Guinier clones, Luisa Avanzo and Dorskamp. Total leaf area, leaf number, leaf area increment and stomatal conductance were evaluated periodically during a 29-day drought period and for 16 days after re-watering. Soil water content and predawn leaf water potential (Psi(wp)) were measured on Days 29 and 45. On the same days, relative water content (RWC), specific leaf area (SLA), nitrogen, chlorophyll, soluble sugars, total phenols, flavanols and antioxidant activity were determined for leaves taken from the bottom to the top of each cutting. Leaves of Luisa Avanzo cuttings grew more rapidly than leaves of Dorskamp and exhibited higher SLA, but lower concentrations of nitrogen, chlorophyll and soluble sugars and lower antioxidant activity per unit area. On Day 29, after withholding water, both clones had closed their stomata, reduced rates of leaf growth, and lower Psi(wp) and RWC; however, the clones differed in their responses to soil water depletion. Compared to Dorskamp, Luisa Avanzo closed its stomata earlier and maintained higher Psi(wp), but lower RWC and leaf sugar concentrations. Antioxidant activity of leaf methanolic extracts decreased in response to water stress only in Luisa Avanzo. Leaf physiology and its modulation by water stress were age dependent in Luisa Avanzo.

Antioxidants↗

Canopy and hydraulic conductance in young, mature and old Douglas-fir trees.

We tested for reductions in water transport with increasing tree size, a key component in determining whether gas exchange and growth are hydraulically limited in tall trees. During the summers of 1998 and 1999, we measured water transport with Granier-type, constant-heat sap flow probes, vapor pressure deficit, and leaf and soil water potentials in overstory Pseudotsuga menziesii (Mirb.) Franco trees in three stands differing in size and age (15, 32 and 60 m in height and about 20, 40 and 450 years in age, respectively) in a P. menziesii-dominated forest in the Pacific Northwest, USA. A total of 24 trees were equipped with sap flow sensors--six 60-m trees, nine 32-m trees and nine 15-m trees. Based on the sap flow measurements and leaf area information estimated from leaf area-sapwood area relationships, we estimated crown-averaged stomatal conductance (GS) and leaf-specific hydraulic conductance (KL). We tested the hypothesis that GS and KL vary inversely with tree height (15 > 32 > 60 m). Analysis of variance of GS ranked as 15 = 60 > 32 m during the early summer and 15 > 60 > 32 m during late season drought. Over the growing season, mean daily GS (+/- SE) was 29.2 +/- 4.4, 24.0 +/- 6.8 and 17.7 +/- 7.2 mmol m-2 s-1 for the 15-, 60- and 32-m trees, respectively. The value of K(L) differed among tree heights only during late season drought and ranked 15 > 32 = 60 m. A hydraulic mass balance suggests that greater sapwood conductivity in 60-m trees compared with 32- and 15-m trees is a likely cause for the departure of the above rankings from those predicted by height and leaf-to-sapwood area ratio.

Circadian Rhythm↗

Temporal variation of microfibril angle in Eucalyptus nitens grown in different irrigation regimes.

In 1990, a 2-ha plantation of Eucalyptus nitens (Deane and Maiden) Maiden was established in southeastern Tasmania and subjected to different irrigation regimes. Point dendrometers were installed in March 1995 to monitor radial stem movement every 15 min over several growing seasons. In this study, data from two growing seasons (1996-1998) were considered. From these measurements, daily increments of stem radius were determined. At the end of the second growing season, we extracted 12-mm cores and measured microfibril angles (MFA) of the wood at high resolution. Microfibril angles were rescaled on a time axis and mapped to daily and distance-based elements. Among treatments, irrigated trees in particular formed higher MFA early in the growing season (September-November) and lower MFA later in the growing season. Trees subjected to cyclic droughts showed clear relationships between MFA and soil water deficits, with MFA increasing in response to water stress release. Increases in MFA were preceded by accelerations in daily increment of stem radius. Among treatments, trees subjected to severe drought had the smallest MFA and generally low fluctuations in MFA. Irrigated trees were susceptible to changes in climate, whereas growth of the trees in the other treatments was limited by water availability. Use of path-analysis showed that temperature had an effect on stem radius increment but not on MFA; wind speed was the only factor that influenced MFA directly. Microfibril angle was correlated with stem shrinking and expansion phases; growth period length and growth rates were positively related to MFA.

Eucalyptus↗

Sap flow of three co-occurring Mediterranean woody species under varying atmospheric and soil water conditions.

We studied the seasonal patterns of water use in three woody species co-occurring in a holm oak forest in northeastern Spain. The three species studied, Quercus ilex L., Phillyrea latifolia L. and Arbutus unedo L., constitute more than 99% of the total basal area of the forest. The study period included the dry seasons of 1999 and 2000. Water use was estimated with Granier-type sap flux sensors. Standard meteorological variables, soil water content and leaf water potentials were also monitored. All monitored individuals reduced leaf-related sap flow (Q(l)) during the summer, concurrent with an increase in soil moisture deficit (SMD). Despite similar maximum Q(l) between species, the decline in Q(l) with increasing SMD was species-dependent. The average reduction in Q(l) between early summer and the peak of the drought was 74% for A. unedo (n = 3), 58% for P. latifolia (n = 3) and 87% for Q. ilex (n = 1). The relationship between canopy stomatal conductance (G(s)) and vapor pressure deficit (D) changed during the course of the drought, with progressively lower G(s) for any given D. Summertime reductions of Q(l) and G(s) were associated with between-species differences in vulnerability to xylem embolism, and with the corresponding degree of native embolism (lowest in P. latifolia and highest in Q. ilex). Our results, combined with previous studies in the same area, outlined differences among the species studied in manner of responding to water shortage, with P. latifolia able to maintain water transport at much lower water potentials than the other two species. In an accompanying experiment, A. unedo responded to an experimental reduction in water availability by reducing Q(l) during the summer. This species also modified its water use between years according to the different seasonal patterns of precipitation. These results are discussed in relation to the possible impacts that climate change will have on Q. ilex-dominated forests.

Ericaceae↗

Interactive effects of temperature and precipitation on soil respiration in a temperate maritime pine forest.

Soil respiration (SR) was monitored periodically throughout 2001 in a Scots pine (Pinus sylvestris L.) stand located in the Belgian Campine region. As expected for a temperate maritime forest, temperature was the dominant control over SR during most of the year. However, during late spring and summer, when soil water content (SWC) was limiting, SR was insensitive to temperature (Q(10) = 1.24). We observed that during prolonged rain-free periods, when SWC was less than 15% (v/v), SR decreased dramatically (up to 50%) and SWC took over control of SR. During such drought periods, however, rain events sometimes stimulated SR and restored temperature control over SR, even though SWC in the mineral soil was low. We hypothesize that restoration of temperature control occurred only when rain events adequately rewetted the uppermost soil layers, where most of the respiratory activity occurred. To quantify the rewetting capacity of rain events, an index (I(w)) was designed that incorporated rainfall intensity, time elapsed since the last rain event, and atmospheric vapor pressure deficit (a proxy for evaporative water losses). To simulate SR fluxes, a model was developed that included the effects of soil temperature and, under drought and non-rewetting conditions (I(w) and SWC < threshold), an SWC response function. The model explained 95% of the temporal variability in SR observed during summer, whereas the temperature function alone explained only 73% of this variability. Our results revealed that, in addition to temperature and SWC, rain plays a role in determining the total amount of carbon released from soils, even in a maritime climate.

Cell Respiration↗

Explanation of vegetation succession in subtropical southern China based on ecophysiological characteristics of plant species.

A stomatal conductance model and a photosynthesis model were applied to field measurements of transpiration and photosynthesis of seven tree species growing in subtropical southern China. Parameter values of drought resistance and tolerance and biochemical assimilation capacity were obtained by means of nonlinear statistical regression, and were used to quantify species succession. The analysis indicated that the models adequately described the ecophysiological behavior of the trees under various environmental conditions. We found a general pattern of decreased drought resistance and tolerance, but increased biochemical assimilation capacity from pines to heliophilus broadleaf trees to mesophilus broadleaf trees. Succession was explained on the basis of these physiological characteristics together with positive feedbacks caused by changes in soil physical properties. The ecophysiological explanation of succession implies that: (1) fitness of a species for a particular succession stage at a particular location can be measured by stomatal behavior and biochemical assimilation capacity under local climate and soil conditions; (2) selection of species for a particular location at a particular succession stage can be guided by the parameter values provided in this study; and (3) succession may be accelerated by selecting trees with large root systems and large soil-root conductances that facilitate soil hydraulic redistribution of water.

China↗

Wood density and anatomy of water-limited eucalypts.

We hypothesized that seedlings grown under water-limited conditions would develop denser wood than seedlings grown under well-watered conditions. Three Eucalyptus species (E. grandis Hill (ex Maiden), E. sideroxylon Cunn. (ex Woolls) and E. occidentalis Endl.) were grown in a temperature-controlled greenhouse for 19 weeks with watering treatments (well-watered and water-limited) applied at six weeks. The water-limitation treatment consisted of four drought cycles. Wood density increased by between 4 and 13% in the water-limited seedlings, but this increase was mainly due to extractive compounds embedded in the cell wall matrix. Once these compounds were removed, the increase was 0-9% and was significant for E. grandis only. Water-limitation significantly reduced mean vessel lumen area; however, this was balanced by a trend toward greater vessel frequency in water-limited plants, and consequently there was no difference in the proportion of stem area allocated to vessels. Conduit efficiency value was lowest in the water-limited plants, indicating that there was a cost in terms of stem hydraulic conductivity for decreasing vessel lumen area. Wood density was negatively correlated with vessel lumen fraction in well-watered plants, but this relationship broke down in the water-limited plants, possibly because of the significantly larger proportion of the stem taken up by pith in water-limited seedlings. Diurnal variation in leaf water potential was positively correlated with wood density in well-watered plants. This relationship did not hold in the water-limited plants owing to the collapse of the pressure gradient between soil and leaf. We conclude that drought periods of greater than 1 month are required to increase wood density in these species and that increases in wood density appear to result in diminished capacity to supply water to leaves.

Dehydration↗