Genotypic variation in drought response of silver birch (Betula pendula): leaf water status and carbon gain

To assess genotypic variation in drought response of silver birch (Betula pendula Roth), we studied the plasticity of 16 physiological traits in response to a 12-14-week summer drought imposed on four clones in two consecutive years. In a common garden experiment, 1-year-old clonal trees from regions with low (550 mm year(-1)) to high rainfall (1270 mm year(-1)) were grown in 45-l pots, and leaf gas exchange parameters, leaf water potentials, leaf osmotic potentials and leaf carbon isotope signatures were repeatedly measured. There were no clonal differences in leaf water potential, but stomatal conductance (gs), net photosynthesis at ambient carbon dioxide concentration, photosynthetic water-use efficiency, leaf carbon isotope composition (delta13C) and leaf osmotic potentials at saturation (Pi0) and at incipient plasmolysis (Pip) were markedly influenced by genotype, especially gs and osmotic adjustment. Genotypes of low-rainfall origin displayed larger osmotic adjustment than genotypes of high-rainfall origin, although their Pi0 and Pip values were similar or higher with ample water supply. Genotypes of low-rainfall origin had higher gs than genotypes of high-rainfall origin under both ample and limited water supply, indicating a higher water consumption that might increase competitiveness in drought-prone habitats. Although most parameters tested were significantly influenced by genotype and treatment, the genotype x treatment interactions were not significant. The genotypes differed in plasticity of the tested parameters and in their apparent adaptation to drought; however, among genotypes, physiological plasticity and drought adaptation were not related to each other. Reduction of gs was the first and most plastic response to drought in all genotypes, and allowed the maintenance of high predawn leaf water potentials during the drought. None of the clones exhibited non-stomatal limitation of photosynthesis. Leaf gs, photosynthetic capacity, magnitude of osmotic adjustment and delta13C were all markedly lower in 2000 than in 1999, indicating root limitation in the containers in the second year.     

Water-use efficiency in cork oak (Quercus suber) is modified by the interaction of water and light availabilities

We studied the interaction of light and water on water-use efficiency in cork oak (Quercus suber L.) seedlings. One-year-old cork oak seedlings were grown in pots in a factorial experiment with four light treatments (68, 50, 15 and 5% of full sunlight) and two irrigation regimes: well watered (WW) and moderate drought stress (WS). Leaf predawn water potential, which was measured at the end of each of two cycles, did not differ among the light treatments. Water-use efficiency, assessed by carbon isotope composition (delta(13)C), tended to increase with increasing irradiance. The trend was similar in the WW and WS treatments, though with lower delta(13)C in all light treatments in the WW irrigation regime. Specific leaf area increased with decreasing irradiance, and was inversely correlated with delta(13)C. Thus, changes in delta(13)C could be explained in part by light-induced modifications in leaf morphology. The relationship between stomatal conductance to water vapor and net photosynthesis on a leaf area basis confirmed that seedlings in higher irradiances maintained a higher rate of carbon uptake at a particular stomatal conductance, implying that shaded seedlings have a lower water-use efficiency that is unrelated to water availability.     

Osmotic regulation in leaves and roots of olive trees during a water deficit and rewatering

We evaluated the osmotic adjustment capacity of leaves and roots of young olive (Olea europaea L.) trees during a period of water deficit and subsequent rewatering. The trials were carried out in Basilicata (40 degrees 24' N, 16 degrees 48' E) on 2-year-old self-rooted olive plants (cv. 'Coratina'). Plants were subjected to one of four drought treatments. After 13 days of drought, plants reached mean predawn leaf water potentials of -0.45 +/- 0.015 MPa (control), -1.65 +/- 0.021 (low stress), -3.25 +/- 0.035 (medium stress) and -5.35 +/- 0.027 MPa (high stress). Total osmotic adjustment increased with increasing severity of drought stress. Trees in the high stress treatment showed total osmotic adjustments ranging between 2.4 MPa at 0500 h and 3.8 MPa at 1800 h on the last day of the drought period. Osmotic adjustment allowed the leaves to reach leaf water potentials of about -7.0 MPa. Active osmotic adjustment at predawn decreased during the rewatering period in both leaves and roots. Stomatal conductance and net photosynthetic rate declined with increasing drought stress. Osmotic adjustment in olive trees was associated with active and passive osmotic regulation of drought tolerance, providing an important mechanism for avoiding water loss.     

Competition for water between walnut seedlings (Juglans regia) and rye grass (Lolium perenne) assessed by carbon isotope discrimination and delta18O enrichment

Container-grown walnut seedlings (Juglans regia L.) were subjected to competition with rye grass (Lolium perenne L.) and to a 2-week soil drying cycle. One and 2 weeks after the beginning of the drought treatment, H2 18O (delta approximately equals +100%) was added to the bottom layer of soil in the plant containers to create a vertical H2 18O gradient. Rye grass competition reduced aboveground and belowground biomass of the walnut seedlings by 60%, whereas drought had no effect. The presence of rye grass reduced the dry weight of walnut roots in the upper soil layer and caused a 50% reduction in lateral root length. Rye grass competition combined with the drought treatment reduced walnut leaf CO2 assimilation rate (A) and leaf conductance (gw) by 20 and 39%, respectively. Transpiration rates in rye grass, both at the leaf level and at the plant or tiller level, were higher than in walnut seedlings. Leaf intrinsic water-use efficiency (A/gw) of walnut seedlings increased in response to drought and no differences were observed between the single-species and mixed-species treatments, as confirmed by leaf carbon isotope discrimination measurements. Measurement of delta18O in soil and in plant xylem sap indicated that the presence of rye grass did not affect the vertical profile of soil water uptake by walnut seedlings. Walnut seedlings and rye grass withdrew water from the top and middle soil layers in well-watered conditions, whereas during the drought treatment, walnut seedlings obtained water from all soil layers, but rye grass took up water from the bottom soil layer only.     

Characteristics of water relations in seedling of Machilus yunnanensis and Cinnamomum camphora under soil drought condition

Introduction Soil drought not only influences agriculture and forestry production, but also brings much trouble for cultivating plants of gardening in the urban. Although the mean annual precipitation of Kunming City, China is 1079 mm, its distribution is…     

Metabolic responses to water deficit in two Eucalyptus globulus clones with contrasting drought sensitivity

We compared the metabolic responses of leaves and roots of two Eucalyptus globulus Labill. clones differing in drought sensitivity to a slowly imposed water deficit. Responses measured included changes in concentrations of soluble and insoluble sugars, proline, total protein and several antioxidant enzymes. In addition to the general decrease in growth caused by water deficit, we observed a decrease in osmotic potential when drought stress became severe. In both clones, the decrease was greater in roots than in leaves, consistent with the observed increases in concentrations of soluble sugars and proline in these organs. In roots of both clones, glutathione reductase activity increased significantly in response to water deficit, suggesting that this enzyme plays a protective role in roots during drought stress by catalyzing the catabolism of reactive oxygen species. Clone CN5 has stress avoidance mechanisms that account for its lower sensitivity to drought compared with Clone ST51.     

Diversity of leaf traits related to productivity in 31 Populus deltoides x Populus nigra clones

To test if some leaf parameters are predictors of productivity in a range of Populus deltoides (Bartr.) Marsh. x P. nigra L. clones, we assessed leaf traits and productivity in 2-month-old rooted cuttings from 31 clones growing in 4-l pots in a greenhouse, under conditions of controlled temperature and optimal irrigation. We evaluated four groups of variables describing (1) productivity (total biomass), (2) leaf growth (total leaf number increment and total leaf area increment rate), (3) leaf structure (specific leaf area and nitrogen and carbon contents) and (4) carbon isotope discrimination (delta), which is negatively correlated with time-integrated water-use efficiency. High-yielding clones did not necessarily display high leaf growth rates, but they displayed a larger total leaf area, lower specific leaf area and lower leaf nitrogen concentration than clones with low productivity. Total leaf area was mainly controlled by maximal individual leaf area and total leaf area increment rate (r = 0.51 and 0.56, respectively). Carbon isotope discrimination did not correlate with total biomass, but it was associated with total number of leaves and total leaf area increment rate (r = 0.39 and 0.45, respectively). Therefore, leaf area and specific leaf area were better indicators of productivity than leaf growth traits. The observed independence of delta from biomass production provides opportunities for selecting poplar clones combining high productivity and high water-use efficiency.     

Evidence for increased sensitivity to nutrient and water stress in a fast-growing hybrid willow compared with a natural willow clone

The hypothesis that fast-growing breeds of willow (Salix spp.) are more sensitive to nutrient and water stress and less efficient in nutrient- and water-use than slower-growing natural willow clones was tested. Cuttings of a natural clone of S. viminalis L. collected in Sweden (L78183) and a hybrid clone of S. schwerinii E. Wolf. x S. viminalis L. ("Tora") were grown outdoors in pots under various experimental conditions in a full-factorial design. The experimental conditions included three fertilization, two irrigation and two temperature regimes. Classical growth analysis techniques, based on an initial and a final harvest, were used as a screening method, together with calculation of intrinsic water-use efficiency (foliar carbon isotope ratio; delta13C). In addition, nitrogen-use efficiency was calculated as the product of nitrogen productivity and mean residence time of nitrogen on an annual basis. There were significant differences in plant structural parameters (leaf area ratio, specific leaf area) and water-use efficiency between the clones. Furthermore, several clone x treatment interaction effects on various growth parameters indicated that the clones adapted to specific environments in different ways. "Tora" plants produced up to 25% more shoot biomass than plants of the natural clone in response to high rates of fertilization and irrigation, whereas clone ranking was reversed in most other treatments. The results support the hypothesis that fast-growing hybrids are more sensitive to nutrient and water stress than slower-growing natural clones. The hypothesis that natural clones have higher resource-use efficiency than fast-growing hybrids was supported with respect to water, but not nitrogen.     

Dry mass allocation, water use efficiency and delta(13)C in clones of Eucalyptus grandis, E. grandis x camaldulensis and E. grandis x nitens grown under two irrigation regimes

Clonal variation in water use efficiency (WUE), dry mass accumulation and allocation, and stable carbon isotope ratio (delta(13)C) of crude leaf fiber extracts was determined in six clones of Eucalyptus grandis W. Hill ex Maiden. grown for 16 months in field lysimeters in two soil water regimes. The relationships between delta(13)C and WUE calculated on the basis of leaf, harvestable stem, shoot and whole-plant dry mass accumulation were investigated. There was no clonal variation in dry mass accumulation but clonal allocation to roots, harvestable stems, branches and leaves varied. Water use efficiencies (mass of plant or plant part/water used over 16 months) differed significantly between clones. The clonal ranking of WUE varied depending on the units of dry mass accumulation used. Significant relationships between delta(13)C values and instantaneous water use efficiencies and ratios of internal leaf to ambient CO(2) concentrations were found only in the high soil water treatment. There were no relationships between delta(13)C values and whole-plant, shoot and harvestable stem water use efficiencies and soil water availability. Values of delta(13)C were negatively correlated with dry mass accumulation in the low soil water treatment. At the whole-plant level, WUE was positively correlated with dry mass accumulation in the high soil water treatment. We found significant differences in delta(13)C values between clones and the clonal rankings in delta(13)C and WUE were maintained in both soil water treatments.     

Carbon isotopic composition, gas exchange, and growth of three populations of ponderosa pine differing in drought tolerance

Effects of water supply on gas exchange, carbon isotopic composition, and relative growth rate were compared among seedlings from three populations of ponderosa pine (Pinus ponderosa Dougl. ex Laws.) grown in a controlled environment chamber. The three populations were chosen to represent high, moderate and low drought tolerance. There was no indication that drought tolerance was related to high water-use efficiency. Populations differed (P < 0.05) in relative growth rate (RGR), but did not differ (P > 0.10) in gas exchange variables or carbon isotope ratio (delta(13)C). Well-watered seedlings had significantly higher RGR, xylem pressure potential (Psi(xpp)), net photosynthesis (A), stomatal conductance to water vapor (g), and lower delta(13)C and instantaneous water-use efficiency than water-stressed seedlings. With decreasing Psi(xpp), A decreased linearly, whereas g decreased exponentially. Seedlings of the highly drought-tolerant population were more sensitive to water availability than seedlings from the other populations; they used water quickly when water was available, but closed their stomata in response to water stress. We conclude that, in ponderosa pine, the drought avoidance mechanism is more important for survival and growth in arid and semiarid environments than the efficient use of water.     

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.     

Clonal variation of Populus tremuloides responses to diurnal drought stress

We have developed an automated microprocessor controlled system for subjecting hydroponically grown plants to drought. Pumps and valves were used to move nutrient solutions into and out of a system of culture vessels in a growth chamber to provide periods of drought. Drought conditions were obtained by exposing the roots of hydroponically grown clones of aspen, Populus tremuloides Michx., to air in culture vessels temporarily emptied of nutrient medium. Over a 3-week period, the daily duration of drought was increased from 0 to 6 h. During this period, the plants became increasingly tolerant to drought, as shown by a decreasing propensity to wilt. All three clones sustained diurnal drought periods of 6 h for up to 5 weeks without detectable deterioration of health. Typical drought stress symptoms were observed including inhibition of growth, increased tissue amino acid content, and decreased water, solute, and turgor potentials in young leaves. In all clones, control plants had leaf water potentials between -1.0 and -1.6 MPa, whereas leaf water potentials of drought-treated plants were significantly lower, ranging from -1.7 to -3.0 MPa. Only one of the clones showed a significant decrease in leaf solute potential in response to drought. The decrease in leaf solute potential paralleled the decrease in water potential resulting in no significant difference in turgor potential. The other two clones had nonsignificant decreases to more negative leaf solute potentials under drought conditions resulting in significantly lowered turgor potentials. Leaf water potentials, solute potentials, and turgor potentials of the drought-treated plants returned to control values within two hours after rewatering. The growth inhibitions observed could not have been the consequence of loss of turgor. These results demonstrate genetic differences among aspen clones in water relations responses to drought.     

锦鸡儿属植物抗旱性研究进展

锦鸡儿属植物是极具开发价值的生态经济型灌木,也是我国西北生态建设的重要植物资源.干旱引起的水分胁迫,对植物的影响主要表现在生长发育及生理代谢过程等方面.文中概述了干旱胁迫条件下的锦鸡儿属植物种子萌发情况,叶片、茎和根的形态结构特征,以及不同程度干旱胁迫对植物组织水分、光合作用、细胞膜系统、渗透调节物质和抗氧化酶系统的影...     

Overexpression of AtNHX5 improves tolerance to both salt and drought stress in Broussonetia papyrifera (L.) Vent

Paper mulberry (Broussonetia papyrifera L. Vent) is well known for its bark fibers, which are used for making paper, cloth, rope, etc. It was found that, in addition to its well-documented role in the enhancement of plant salt tolerance, overexpression of the Na+/H+ antiporter (AtNHX5) gene in paper mulberry plants showed high drought tolerance. After exposure to water deficiency and salt stress, the wild-type (WT) plants all died, while the AtNHX5-overexpressing plants remained alive under high salt stress, and had a higher survival rate (>66%) under drought stress. Measurements of ion levels indicated that Na+ and K+ contents were all higher in AtNHX5-overexpressing leaves than in WT leaves in high saline conditions. The AtNHX5 plants had higher leaf water content and leaf chlorophyll contents, accumulated more proline and soluble sugars, and had less membrane damage than the WT plants under water deficiency and high saline conditions. Taken together, the results indicate that the AtNHX5 gene could enhance the tolerance of paper mulberry plants to multiple environmental stresses by promoting the accumulation of more effective osmolytes (ions, soluble sugars, proline) to counter the osmotic stress caused by abiotic factors.     

Modeling photosynthesis in olive leaves under drought conditions

We quantified parameters for a model of leaf-level photosynthesis for olive, and tested the model against an independent dataset. Specific temperature-dependence parameters of the model for olive leaves were measured, as well as the relationship of the model parameters with area-based leaf nitrogen (N) content. The effect of soil water deficit on leaf photosynthesis was examined by applying two irrigation treatments to 29-year-old trees growing in a plantation: drip irrigation sufficient to meet the crop water requirements (I) and dry-farming (D). In both treatments, leaves had a higher photosynthetic capacity in April than in August. In August, photosynthetic capacity was lower in D trees than in I trees. Leaf photosynthetic capacity was linearly and positively related to leaf N content on an area basis (N(a)) and to leaf mass per unit area (LMA), and the regression slope varied with irrigation treatment. The seasonal reduction in N(a) was used in the model to predict photosynthesis under drought conditions. Olive leaves showed a clear limitation of photosynthesis by triose phosphate utilization (TPU) even at 40 degrees C, and the data suggest that olive invests fewer resources in TPU than other species. The seasonal decrease in photosynthetic capacity moderated the stomatal limitation to carbon dioxide (CO(2)) fixation as soil water deficit increased. Further, it enabled leaves to operate close to the transition point between photosynthetic limitation due to RuBP carboxylation capacity and that due to RuBP regeneration capacity, and resulted in a near constant value of internal CO(2) concentration from April to August. Under well watered conditions, N-use efficiency of the olive leaves was enhanced at the expense of reduced water-use efficiency.     

Variation in drought resistance, drought acclimation and water conservation in four willow cultivars used for biomass production

Growth and water-use parameters of four willow (Salix spp.) clones grown in a moderate drought regime or with ample water supply were determined to characterize their water-use efficiency, drought resistance and capacity for drought acclimation. At the end of the 10-week, outdoor pot experiment, clonal differences were observed in: (1) water-use efficiency of aboveground biomass production (WUE); (2) resistance to xylem cavitation; and (3) stomatal conductance to leaf-specific, whole-plant hydraulic conductance ratio (g(st)/K(P); an indicator of water balance). Across clones and regimes, WUE was positively correlated with the assimilation rate to stomatal conductance ratio (A/g(st)), a measure of instantaneous water-use efficiency. Both of these water-use efficiency indicators were generally higher in drought-treated trees compared with well-watered trees. However, the between-treatment differences in (shoot-based) WUE were smaller than expected, considering the differences in A/g(st) for two of the clones, possibly because plants reallocated dry mass from shoots to roots when subject to drought. Higher root hydraulic conductance to shoot hydraulic conductance ratios (K(R)/K(S)) during drought supports this hypothesis. The same clones were also the most sensitive to xylem cavitation and, accordingly, showed the strongest reduction in g(st)/K(P) in response to drought. Drought acclimation was manifested in decreased g(st), g(st)/K(P), osmotic potential and leaf area to vessel internal cross-sectional area ratio, and increased K(R), K(P) and WUE. Increased resistance to stem xylem cavitation in response to drought was observed in only one clone. It is concluded that WUE and drought resistance traits are inter-linked and that both may be enhanced by selection and breeding.     

Growth, leaf morphology, water use and tissue water relations of Eucalyptus globulus clones in response to water deficit

Changes in leaf size, specific leaf area (SLA), transpiration and tissue water relations were studied in leaves of rooted cuttings of selected clones of Eucalyptus globulus Labill. subjected to well-watered or drought conditions in a greenhouse. Significant differences between clones were found in leaf expansion and transpiration. There was a significant clone x treatment interaction on SLA. Water stress significantly reduced osmotic potential at the turgor loss point (Pi0) and at full turgor (Pi100), and significantly increased relative water content at the turgor loss point and maximum bulk elastic modulus. Differences in tissue water relations between clones were significant only in the mild drought treatment. Among clones in the drought treatments, the highest leaf expansion and the highest increase in transpiration during the experiment were measured in those clones that showed an early and large decrease in Pi0 and Pi100.     

Contrasting physiological responses of two co-occurring eucalypts to seasonal drought at restored bauxite mine sites

This study describes the physiological response of two co-occurring tree species (Eucalyptus marginata and Corymbia calophylla) to seasonal drought at low- and high-quality restored bauxite mine sites in south-western Australia. Seasonal changes in photosynthesis (A), stomatal conductance (g(s)), leaf water potential (ψ), leaf osmotic potential (ψ), leaf relative water content (RWC) and pressure-volume analysis were captured over an 18-month field study to (i) determine the nature and severity of physiological stress in relation to site quality and (ii) identify any physiological differences between the two species. Root system restriction at the low-quality site reduced maximum rates of gas exchange (g(s) and A) and increased water stress (midday ψ and daily RWC) in both species during drought. Both species showed high stomatal sensitivity during drought; however, E. marginata demonstrated a higher dehydration tolerance where ψ and RWC fell to -3.2 MPa and 73% compared with -2.4 MPa and 80% for C. calophylla. Corymbia calophylla showed lower g(s) and higher ψ and RWC during drought, indicating higher drought tolerance. Pressure-volume curves showed that cell-wall elasticity of E. marginata leaves increased in response to drought, while C. calophylla leaves showed lower osmotic potential at zero turgor in summer than in winter, indicating osmotic adjustment. Both species are clearly able to tolerate seasonal drought at hostile sites; however, by C. calophylla closing stomata earlier in the drought cycle, maintaining a higher water status during drought and having the additional mechanism of osmotic adjustment, it may have a greater capacity to survive extended periods of drought.     

花吊丝竹对干旱胁迫的光合和生理响应

通过盆栽实验,研究了花吊丝竹在自然耗水的干旱胁迫处理下土壤含水量、叶片水势、叶片光合作用参数、叶绿素荧光参数以及相关酶类生理指标的变化。结果表明:(1)干旱胁迫下,土壤含水量与叶片水势显著下降,叶片的净光合速率、气孔导度和蒸腾速率均有不同程度下降,胞间CO2浓度呈先降后升的趋势,而气孔限制值则呈先升后降的趋势,说明轻度干旱胁迫下,气孔限制是花吊丝竹净光合速率降低的主要因素,重度干旱胁迫下,非气孔限制是净光合速率降低的主要因素。(2)花吊丝竹叶片光系统Ⅱ的实际光化学效率、表观光合电子传递速率和非光化学猝灭系数在胁迫后段(15~30 d)呈显著下降趋势,而叶片初始荧光在后期呈上升趋势,表明随胁迫程度加重,PSⅡ的结构受到较严重的损伤。(3)随干旱时间的延长,脯氨酸和丙二醛的含量均显著上升,而超氧化物歧化酶和过氧化氢酶的活性则先升后降,这表明随胁迫程度加深花吊丝竹细胞结构被破坏。综上所述,干旱胁迫环境下,花吊丝竹叶片光合作用和保护酶类均发生相应的变化,其自身能够提高叶片对光能的捕获能力、提升光能转化效率、增强叶片中的酶活性以及减少热能的耗散等形式,实现对干旱胁迫较强的忍耐性和较好的适应性。     

Drought resistance of two hybrid Populus clones grown in a large-scale plantation

Poplar hybrids were grown with irrigation in a large-scale plantation to investigate the mechanisms underlying clonal differences in drought resistance. Beginning in spring 1992, Populus trichocarpa x P. deltoides (TD) and P. deltoides x P. nigra (DN) cuttings received 46, 76, or 137 cm year(-1) of irrigation to supplement the 18-20 cm of annual precipitation, and all trees received the same fertilization regime. Stem volume, assessed as the square of stem diameter at breast height times tree height (D(2)H), and water relations of the trees were studied from the end of their second growing season until the end of their fifth growing season. By the end of the second growing season, stem volume of Clone TD was 40-146% larger than that of Clone DN, but stem volume growth was independent of irrigation in excess of 46 cm year(-1) in both clones. During the third growing season, stem volume growth of both clones was limited by both the 46- and 76-cm irrigation treatments, so that by the end of the third growing season trees in the 46-cm irrigation treatment were only half the size of trees in the 137-cm irrigation treatment. These treatment differences were maintained through the fifth growing season. Although stem volumes of Clone TD trees in the 76- and 137-cm irrigation treatments were larger than the corresponding values for Clone DN trees at the end of the third growing season (1994), these clonal differences gradually decreased in subsequent years and were not detectable after 5 years, because stem volume relative growth rate of Clone DN was greater than that of Clone TD in all treatments. Although both clones exhibited similar predawn leaf water potentials, Clone DN typically maintained higher midday leaf water potentials, suggesting better stomatal control of water loss. Clonal and treatment differences in osmotic potential at full turgor were minimal and could not explain the clonal differences in drought resistance. Root density and root density to stem volume ratio increased more in response to moderate drought in Clone DN than in Clone TD, resulting in enhanced drought resistance (high stem volume growth rate under moderate drought conditions) and an increased capacity to withdraw water from the soil. We conclude that the greater drought resistance of Clone DN compared with Clone TD was the result of the maintenance of a more favorable water balance by stomatal regulation and greater carbon allocation to roots during the early stages of drought. However, the low root density to stem volume ratio in Clone DN growing in the 46-cm irrigation treatment suggests that severe water limitation restricted the preferential allocation of carbon to belowground tissues, so that both root and shoot growth were constrained by severe drought.