Xylem vulnerability to cavitation in Pseudotsuga menziesii and Pinus ponderosa from contrasting habitats

In the Rocky Mountains, ponderosa pine (Pinus ponderosa (ssp.) ponderosa Dougl. ex P. Laws. & C. Laws) often co-occurs with Douglas-fir (Pseudotsuga menziesii var. glauca (Mayr) Franco). Despite previous reports showing higher shoot vulnerability to water-stress-induced cavitation in ponderosa pine, this species extends into drier habitats than Douglas-fir. We examined: (1) whether roots and shoots of ponderosa pine in riparian and slope habitats are more vulnerable to water-stress-induced cavitation than those of Douglas-fir; (2) whether species-specific differences in vulnerability translate into differences in specific conductivity in the field; and (3) whether the ability of ponderosa pine to extend into drier sites is a result of (a) greater plasticity in hydraulic properties or (b) functional or structural adjustments. Roots and shoots of ponderosa pine were significantly more vulnerable to water-stress-induced cavitation (overall mean cavitation pressure, Psi(50%) +/- SE = -3.11 +/- 0.32 MPa for shoots and -0.99 +/- 0.16 MPa for roots) than those of Douglas-fir (Psi(50%) +/- SE = -4.83 +/- 0.40 MPa for shoots and -2.12 +/- 0.35 MPa for roots). However, shoot specific conductivity did not differ between species in the field. For both species, roots were more vulnerable to cavitation than shoots. Overall, changes in vulnerability from riparian to slope habitats were small for both species. Greater declines in stomatal conductance as the summer proceeded, combined with higher allocation to sapwood and greater sapwood water storage, appeared to contribute to the ability of ponderosa pine to thrive in dry habitats despite relatively high vulnerability to water-stress-induced cavitation.     

Xylem vulnerability to cavitation varies among poplar and willow clones and correlates with yield

Xylem vulnerability to cavitation is a promising criterion for identifying trees with high drought tolerance, but traditional techniques for measuring cavitation resistance are unsuitable for screening large numbers of genotypes. We tested the potential of the new Cavitron technique for high throughput screening of cavitation resistance in five poplar (Populus spp.) and four willow (Salix spp.) clones. The Cavitron technique enabled the screening of three to four clones per day with sufficient accuracy to reveal significant differences between clones. Because intraspecific screening may be better carried out through the identification of correlated and more easily measured traits, we attempted to identify accessible parameters that correlate to cavitation resistance. Variability in vulnerability to cavitation across clones was poorly correlated with anatomical traits such as vessel diameter, vessel wall strength, wood density and fiber wall thickness; however, a striking correlation was established between cavitation resistance and aboveground biomass production, indicating a possible trade-off between xylem safety and growth potential.     

Introduction of ponderosa pine and Douglas-fir to Argentina

Patterns of shoot elongation of 2-yr seedlings from native North American populations of ponderosa pine and Douglas-fir were compared to those of Argentine land races originating from unknown provenances. The comparisons were conducted in Moscow, Idaho (USA), and suggested that the ponderosa pine land race was descended from a California provenance at low or middle elevations but that the growth potential of the land race was only mediocre in comparison to eight native populations. The Douglas-fir land race exhibited a relatively high growth potential in comparison to 19 native provenances and undoubtedly originated from a mild coastal environment. The results provide concrete recommendations for upgrading the growth potential of the Argentine land races by importing germ plasm of specific provenances.     

Effects of summer drought and winter freezing on stem hydraulic conductivity of Rhododendron species from contrasting climates

We studied the limits to maximum water transport in three diffuse-porous evergreen shrubs exposed to frequent winter freeze-thaw events (Rhododendron maximum L. and R. catawbiense Michaux from the Appalachian Mountains) and to a severe summer drought (R. macrophyllum G. Don. from the Oregon Cascades). Percent loss of hydraulic conductivity (PLC), vulnerability curves to xylem embolism and freezing point temperatures of stems were measured over 2 years. Controlled freeze-thaw experiments were also conducted to determine the effect of thaw rate on PLC. During both years, native PLC was significantly higher in winter than in summer for R. macrophyllum. Seasonal changes in PLC were variable in both R. catawbiense and R. maximum. Only R. maximum plants growing in gaps or clearings showed higher PLC than understory plants. A rapid (2-4 day) natural recovery of high native PLC during the winter was observed in both R. maximum and R. macrophyllum. Compared with the bench-dehydration method, vulnerability curves based on the air-injection method consistently had less negative slopes and greater variation. Fifty percent PLC (PLC(50)) obtained from vulnerability curves based on the dehydration method occurred at -1.75, -2.42 and -2.96 MPa for R. catawbiense, R. maximum and R. macrophyllum, respectively. Among the study species, R. macrophyllum, which commonly experiences a summer drought, had the most negative water potential at PLC(50). In all species, stem freezing point temperatures were not consistently lower in winter than in summer. A single fast freeze-thaw event significantly increased PLC, and R. catawbiense had the highest PLC in response to freezing treatments. Recovery to control PLC values occurred if a low positive hydraulic pressure was maintained during thawing. Rhododendron macrophyllum plants, which commonly experience few freeze-thaw events, had large stem diameters, whereas plants of R. catawbiense, which had small stem diameters, suffered high embolism in response to a single freeze-thaw event. Both drought-induced and winter-induced embolism caused a significant reduction in hydraulic conductivity in all species during periods when drought or freeze-thaw events occurred in their native habitats. However, rapid recovery of PLC following freezing or drought maintained the species above their relatively low margins of safety for complete xylem dysfunction.     

Limited response of ponderosa pine bole defenses to wounding and fungi

Tree defense against bark beetles (Curculionidae: Scolytinae) and their associated fungi generally comprises some combination of constitutive (primary) and induced (secondary) defenses. In pines, the primary constitutive defense against bark beetles consists of preformed resin stored in resin ducts. Induced defenses at the wound site (point of beetle entry) in pines may consist of an increase in resin flow and necrotic lesion formation. The quantity and quality of both induced and constitutive defenses can vary by species and season. The inducible defense response in ponderosa pine is not well understood. Our study examined the inducible defense response in ponderosa pine using traumatic mechanical wounding, and wounding with and without fungal inoculations with two different bark beetle-associated fungi (Ophiostoma minus and Grosmannia clavigera). Resin flow did not significantly increase in response to any treatment. In addition, necrotic lesion formation on the bole after fungal inoculation was minimal. Stand thinning, which has been shown to increase water availability, had no, or inconsistent, effects on inducible tree defense. Our results suggest that ponderosa pine bole defense against bark beetles and their associated fungi is primarily constitutive and not induced.     

Water limitations to carbon exchange in old-growth and young ponderosa pine stands

We investigated the impact of seasonal soil water deficit on the processes driving net ecosystem exchange of carbon (NEE) in old-growth and recently regenerating ponderosa pine (Pinus ponderosa Doug. ex Laws.) stands in Oregon. We measured seasonal patterns of transpiration, canopy conductance and NEE, as well as soil water, soil temperature and soil respiration. The old-growth stand (O) included two primary age classes (50 and 250 years), had a leaf area index (LAI) of 2.1 and had never been logged. The recently regenerating stand (Y) consisted predominantly of 14-year-old ponderosa pine with an LAI of 1.0. Both stands experienced similar meteorological conditions with moderately cold wet winters and hot dry summers. By August, soil volumetric water content within the upper 30 cm had declined to a seasonal minimum of 0.07 at both sites. Between April and June, both stands showed similar rates of transpiration peaking at 0.96 mm day(-1); thereafter, trees at the Y site showed increasing drought stress with canopy stomatal resistance increasing 6-fold by mid-August relative to values for trees at the O site. Over the same period, predawn water potential (psi(pd)) of trees at the Y site declined from -0.54 to -1.24 MPa, whereas psi(pd) of trees at the O site remained greater than -0.8 MPa throughout the season. Soil respiration at the O site showed a strong seasonal correlation with soil temperature with no discernible constraints imposed by declining soil water. In contrast, soil respiration at the Y site peaked before seasonal maximal soil temperatures and declined thereafter with declining soil water. No pronounced seasonal pattern in daytime NEE was observed at either site between April and September. At the Y site this behavior was driven by concurrent soil water limitations on soil respiration and assimilation, whereas there was no evidence of seasonal soil water limitations on either process at the O site.     

Pricing ecosystem resilience in frequent-fire ponderosa pine forests

Dry forests across the United States have become subject to declining resilience and, consequently, increased vulnerability to catastrophic wildfires. These disturbances cause severe environmental and social damages and may dislodge the forest into a different ecological regime. Forests provide many valuable services, such as the provisioning of timber and the sequestering of carbon that would otherwise contribute to climate change. The high-severity conflagrations that have become regular occurrences in many dry forests impinge the delivery of such benefits, particularly in the event of a regime shift. Sustainable forest management should take these risks into account. This article analyzes the economics of resilience in dry forests with respect to catastrophic fires and ecological thresholds. We illustrate how to price ecosystem resilience for the fire-prone ponderosa pine forests of the western United States. This analysis demonstrates that pricing forest resilience also establishes the economic value of ecological restoration with respect to ecosystem services, thereby operationalizing forest management as an investment in natural capital.     

Changes in physiological attributes of ponderosa pine from seedling to mature tree

Plant physiological models are generally parameterized from many different sources of data, including chamber experiments and plantations, from seedlings to mature trees. We obtained a comprehensive data set for a natural stand of ponderosa pine (Pinus ponderosa Laws.) and used these data to parameterize the physiologically based model, TREGRO. Representative trees of each of five tree age classes were selected based on population means of morphological, physiological, and nearest neighbor attributes. Differences in key physiological attributes (gas exchange, needle chemistry, elongation growth, needle retention) among the tree age classes were tested. Whole-tree biomass and allocation were determined for seedlings, saplings, and pole-sized trees. Seasonal maxima and minima of gas exchange were similar across all tree age classes. Seasonal minima and a shift to more efficient water use were reached one month earlier in seedlings than in older trees because of decreased soil water availability in the rooting zone of the seedlings. However, carbon isotopic discrimination of needle cellulose indicated increased water-use efficiency with increasing tree age. Seedlings had the lowest needle and branch elongation biomass growth. The amount of needle elongation growth was highest for mature trees and amount of branch elongation growth was highest for saplings. Seedlings had the highest biomass allocation to roots, saplings had the highest allocation to foliage, and pole-sized trees had the highest allocation to woody tissues. Seedlings differed significantly from pole-sized and older trees in most of the physiological traits tested. Predicted changes in biomass with tree age, simulated with the model TREGRO, closely matched those of trees in a natural stand to 30 years of age.     

Xylem cavitation in roots and stems of Douglas-fir and white fir

Roots of hardwoods have been shown to be more vulnerable to xylem cavitation than stems. This study examined whether this pattern is also observed in a conifer species. Vulnerability to cavitation was determined from the pressure required to inject air into the vascular system of hydrated roots and stems, and reduce hydraulic conductance of the xylem. According to the air-seeding hypothesis for the cavitation mechanism, these air pressures predict the negative xylem pressure causing cavitation in dehydrating stems. This was evaluated for stems of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) and white fir (Abies concolor (Gord. & Glend.) Lindl.). The air-injection method was applied to roots and stems of different sizes and positions in Douglas-fir trees. Roots, especially smaller roots with a xylem diameter < 5 mm, were more vulnerable to cavitation than stems. Mean cavitation pressure for smaller roots was -2.09 +/- 0.42 versus -3.80 +/- 0.19 MPa for larger roots (> 8 mm diameter). Within the shoot system, smaller stems (< 5 mm diameter) were most vulnerable to cavitation, having a mean cavitation pressure of -4.23 +/- 0.565 versus -5.27 +/- 0.513 MPa for large stems (> 8 mm diameter). There was no correlation between tracheid diameter and mean cavitation pressure within root or stem systems, despite larger tracheid diameters in roots (23.3 +/- 3.9 micro m) than in stems (9.2 +/- 1.6 micro m). Smaller safety margins from cavitation in roots may be beneficial in limiting water use during mild drought, and in protecting the stem from low xylem pressures during extreme drought.     

Estimating understory vegetation response to multi-nutrient fertilization in Douglas-fir and ponderosa pine stands

Models were developed to predict understory vegetation response to multi-nutrient fertilization at six conifer-forested stands in the inland Northwest United States. Equations are presented to estimate how fertilization as well as other factors impacting understory production in the inland Northwest change total understory vegetation production and the production of three individual lifeforms (shrubs, forbs, and grasses and grass-likes). Overstory stand density was found to have the greatest impact on understory production, and regardless of factors such as fertilization or precipitation, large stand densities will limit understory production. At lower stand densities, multi-nutrient fertilization as well as greater amounts of precipitation will increase understory production. These factors were also found to be synergistic; thus, greater amounts of precipitation increase the effects of multi-nutrient fertilization on understory production. For sites of the same stand density, Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] was shown to have a greater negative impact on understory production than ponderosa pine (Pinus ponderosa Dougl. ex Laws.). The models predict that multi-nutrient fertilization of ponderosa pine stands will produce increases in understory production across a broader range of stand densities.     

Recruitment of ponderosa pine seedlings in the Cascade Range

Abiotic stresses on seedling regeneration in xeric ecosystems are great, hence recruitment processes can be facilitated by stand factors that ameliorate the germinant-scale microenvironment. An experiment was conducted on the eastern slope of the Cascade Range to test the effects of shrub cover, simulated seed caching, and substrate on the recruitment of Pacific ponderosa pine (Pinus ponderosa var. ponderosa Dougl. ex Laws.) seedlings. Failure rates of seeds sowed in exclosures were large, with less than 30% emerging as germinants in the spring following fall sowing. Simulated seed caching improved emergence rates by more than sevenfold and was responsible for 88% of all spring germinants. Emergence rates were lowest from uncached seeds on litter. Just 16% of the crop survived the summer and fall to the month of November, or less than 5 months after emergence. Shrub cover did not affect emergence rates, but establishment rates were higher: seedlings beneath shrubs succumbed to desiccation at a slower rate than unshaded seedlings. By August there were 2.3 times more survivors at shrub-shaded sites than unshaded sites, and by the end of fall, when seedlings were considered established, more than 78% existed beneath shrubs. This study provides evidence that the natural recruitment of ponderosa pine seedlings is facilitated by the occurrence of the species’ common shrub associates.     

Accentuation of gas exchange gradients in flushes of ponderosa pine exposed to ozone

Two-year-old ponderosa pine seedlings (Pinus ponderosa Laws.) were exposed to episodic O(3) concentrations in open-top chambers for two consecutive growing seasons (June through September of 1990 and 1991). Near the end of the second season of O(3) exposure, gas exchange was measured on needles of surviving flushes at saturating CO(2) and photosynthetic photon flux density (PPFD). Both photosynthetic capacity (A(sat)) and stomatal conductance to water vapor (g(wv)) declined linearly with needle age but differences within a flush were also found. Gas exchange rates of needles from the base of the current-year flush were significantly lower than those of needles from the top of the flush, even though age differences between needles were negligible. Although most measurements were conducted at saturating CO(2), similar patterns of gas exchange were also found at 350 micro mol mol(-1) CO(2), indicating that photosynthesis of needles at the bottom of the flush was more strongly affected by O(3) than that of needles at the top of the flush, even though the potential for O(3) uptake was probably less in needles at the bottom of the flush because of reduced stomatal conductance. Carboxylation efficiency (deltaA/deltaC(i)), the linear slope of the A/C(i) response, was highly correlated with A(sat), varying with needle age, needle position in the flush and O(3) exposure, but the magnitude of the reductions was greater than for A(sat). We conclude that susceptibility to O(3) damage among needles of an individual seedling varies not only with needle age but also with needle position, and that reductions in photosynthetic capacity may not be directly attributable to increased uptake of the pollutant. The data also indicate the need to consider within-flush variation when estimating whole-plant carbon gain and O(3) uptake.     

Seed rain of ponderosa pine beneath partial overstories

An analysis was conducted of the 1999 ponderosa pine (Pinus ponderosa var. ponderosa Dougl. ex Laws.) seed crop at two stands of differing overstory density in each of two sites that differed in productivity (forest community series) on the east slope of the Cascade Range in central Oregon, USA. A total 2,166 viable seeds were collected between 2 September 1999 and 6 June 2000. Total seed quantities varied more by site than by tree density, with the xeric site producing six-fold greater seed yields. Within each site, the stand of higher density produced more viable seed. Per-tree cone yields were also greater at the site of lesser productivity. At three of the four stands, tree fecundity (as measured by cone counts) was positively related to diameter at breast height and height, but not to live crown ratio. A difference in temporal seedfall patterns among the two sites occurred late and was relatively minor: most seedfall occurred within the first month after it started regardless of absolute seed production. Seedfall distribution varied spatially within stands, but no seed trap (of 55) received less than an equivalent 28 thousand seeds per hectare. Seedfall was not perceived to be a limiting factor on natural regeneration at any of the four partial-overstory stands in this study.     

Xylem cavitation and loss of hydraulic conductance in western hemlock following planting

Following planting, western hemlock (Tsuga heterophylla (Raf.) Sarg.) seedlings experience water stress and declining xylem pressure potential (Psi(x)). Low Psi(x) can result in xylem cavitation and embolism formation, causing a decline in hydraulic conductance. This study focused on the relationship between Psi(x), xylem cavitation and transpiration (E) of newly planted seedlings. Leaf specific hydraulic conductance (k(AB)) declined from 0.56 to 0.09 mmol m(-2) s(-1) MPa(-1) over a 9-day period. Stomatal conductance (g(s)) declined from 143.5 to 39.15 mmol m(-2) s(-1) over the same period without an associated change in environmental conditions. A vulnerability profile indicated a 30% loss in hydraulic conductivity when seedlings experienced a Psi(x) between -2.5 and -3.0 MPa. A Psi(x) of -4.0 MPa led to a complete loss of conductivity. We conclude that following planting, western hemlock seedlings often experience Psi(x) values that are low enough to cause xylem cavitation and a decline in k(AB).     

Nutrient availability alters belowground respiration of ozone-exposed ponderosa pine

Exposure to ozone (O(3)) and changes in soil fertility influence both the metabolism of plant roots and their interaction with rhizosphere organisms. Because one indication of altered root metabolism is a change in belowground respiratory activity, we used specially designed measurement chambers to assess the effects of O(3) and nutrient availability on belowground respiratory activity of potted three-year-old ponderosa pine (Pinus ponderosa Dougl. ex Laws.). Seedlings were exposed to a factorial combination of three O(3) treatments and three fertilization treatments in open-top O(3) exposure chambers. Ozone exposure decreased and high nutrient supply increased total plant dry weight, but root/shoot ratios were not affected. In general, exposure to O(3) increased rates of belowground O(2) uptake and CO(2) release and the respiratory quotient (RQ, CO(2)/O(2)), although seasonal differences were detected. In October, following the second season of O(3) exposure, rates of belowground O(2) uptake and CO(2) release and RQ were increased in trees in the high-O(3) exposure treatment by 22, 73 and 32%, respectively, over values in control trees in charcoal-filtered air. Increasing nutrient supply resulted in decreasing rates of belowground O(2) uptake and CO(2) release but it had little effect on RQ. In the high-nutrient supply treatment, rates of belowground O(2) uptake and CO(2) release were decreased by 38 and 39%, respectively, compared with rates in the low-nutrient supply treatment. At the end of the second growing season, the high-nutrient supply treatment had decreased lateral root total nonstructural carbohydrates by 22% compared with the low-nutrient supply treatment. Nutrient availability altered the belowground respiratory response to O(3), such that the response to O(3) was greatest in the low-nutrient supply treatment. Significant O(3) effects on belowground respiratory activity were apparent before any reduction in total plant growth was found, suggesting that roots and rhizosphere organisms may be early indicators of physiological dysfunction in stressed seedlings.     

Response of antelope bitterbrush to repeated prescribed burning in Central Oregon ponderosa pine forests

Antelope bitterbrush is a dominant shrub in many interior ponderosa pine forests in the western United States. How it responds to prescribed fire is not well understood, yet is of considerable concern to wildlife and fire managers alike given its importance as a browse species and as a ladder fuel in these fire-prone forests. We quantified bitterbrush cover, density, and biomass in response to repeated burning in thinned ponderosa pine forests. Low- to moderate-intensity spring burning killed the majority of bitterbrush plants on replicate plots. Moderately rapid recovery of bitterbrush density and cover resulted from seedling recruitment plus limited basal sprouting. Repeated burning after 11 years impeded the recovery of the bitterbrush community. Post-fire seed germination following the repeated burns was 3–14-fold lower compared to the germination rate after the initial burns, while basal sprouting remained fairly minor. After 15 years, bitterbrush cover was 75–92% lower on repeated-burned compared to unburned plots. Only where localized tree mortality resulted in an open stand was bitterbrush recovery robust. By controlling bitterbrush abundance, repeated burning eliminated the potential for wildfire spread when simulated using a customized fire behavior model. The results suggest that repeated burning is a successful method to reduce the long-term fire risk imposed by bitterbrush as an understory ladder fuel in thinned pine stands. Balancing the need to limit fire risk yet provide adequate bitterbrush habitat for wildlife browse will likely require a mosaic pattern of burning at the landscape scale or a burning frequency well beyond 11 years to allow a bitterbrush seed crop to develop.     

Shoot and root vulnerability to xylem cavitation in four populations of Douglas-fir seedlings

The objectives of this study were to assess the range of genotypic variation in the vulnerability of the shoot and root xylem of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) seedlings to water-stress-induced cavitation, and to assess the trade-off between vulnerability to cavitation and conductivity per unit of stem cross-sectional area (k(s)), both within a species and within an individual tree. Douglas-fir occupies a broad range of environments and exhibits considerable genetic variation for growth, morphology, and drought hardiness. We chose two populations from each of two varieties (the coastal var. menziesii and the interior var. glauca) to represent environmental extremes of the species. Vulnerability curves were constructed for shoots and roots by plotting the percentage loss in conductivity versus water potential. Vulnerability in shoot and root xylem varied genetically with source climate. Stem xylem differed in vulnerability to cavitation between populations; the most mesic population, coastal wet (CW), was the most susceptible of the four populations. In the roots, the most vulnerable population was again CW; the interior wet (IW) population was moderately susceptible compared with the two dry populations, coastal dry (CD) and interior dry (ID). Root xylem was more susceptible to cavitation than stem xylem and had significantly greater k(s). The trade-off between vulnerability to cavitation and k(s), however, was not evident across populations. The most vulnerable population (CW) had a shoot k(s) of 0.534 +/- 0.067 &mgr;mol m(-2) s(-1) MPa(-1), compared with 0.734 +/- 0.067 &mgr;mol m(-2) s(-1) MPa(-1) for the less vulnerable CD stems. In the roots, IW was more vulnerable than ID, but had the same k(s).     

Large mulches and manual release enhance growth of ponderosa pine seedlings

Woody and herbaceous vegetation that captures scarce soil moisture often kills or stunts the growth of conifer seedlings in young plantations in the Sierra Nevada of California. Two methods for excluding this vegetation are to apply large (3-m×3-m), long-lasting (at least 5 years) mulches around the seedlings soon after planting, or to repeatedly grub seedlings of competing vegetation in study plots. Both techniques were tested and proved to be effective, significantly increasing ponderosa pine diameter and height relative to the control and to seedlings surrounded by small paper mulches. The effective treatments were large enough to enable conifer seedlings to capture site resources unimpeded by competing plants. Eight species of shrubs, seventeen forbs, four grasses, and one fern grew naturally on the study site.Results suggest that the forester now has two biologically effective means for controlling unwanted vegetation and for attaining rapid early pine growth in environments similar to the study area. Although more expensive, the large heavy polyester mulches ($9.90 per seedling over 5 years) give the forester an alternative to manual release ($2.05 per seedling over 5 years).     

Use of a simulation model and ecosystem flux data to examine carbon-water interactions in ponderosa pine

Drought stress plays an important role in determining both the structure and function of forest ecosystems, because of the close association between the carbon (C) and hydrological cycles. We used a detailed model of the soil-plant-atmosphere continuum to investigate the links between carbon uptake and the hydrological cycle in a mature, open stand of ponderosa pine (Pinus ponderosa Dougl. ex Laws.) at the Metolius river in eastern Oregon over a 2-year period (1996-1997). The model was parameterized from local measurements of vegetation structure, soil properties and meteorology, and tested against independent measurements of ecosystem latent energy (LE) and carbon fluxes and soil water content. Although the 2 years had very different precipitation regimes, annual uptake of C and total transpiration were similar in both years, according to both direct observation and simulations. There were important differences in ratios of evaporation to transpiration, and in the patterns of water abstraction from the soil profile, depending on the frequency of summer storms. Simulations showed that, during periods of maximum water limitation in late summer, plants maintained a remarkably constant evapotranspirative flux because of deep rooting, whereas changes in rates of C accumulation were determined by interactions between atmospheric vapor pressure deficit and stomatal conductance. Sensitivity analyses with the model suggest a highly conservative allocation strategy in the vegetation, focused belowground on accessing a soil volume large enough to buffer summer droughts, and optimized to account for interannual variability in precipitation. The model suggests that increased allocation to leaf area would greatly increase productivity, but with the associated risk of greater soil water depletion and drought stress in some years. By constructing sparse canopies and deep rooting systems, these stands balance reduced productivity in the short term with risk avoidance over the long term.