Hydraulic redistribution of soil water during summer drought in two contrasting Pacific Northwest coniferous forests

The magnitude of hydraulic redistribution of soil water by roots and its impact on soil water balance were estimated by monitoring time courses of soil water status at multiple depths and root sap flow under drought conditions in a dry ponderosa pine (Pinus ponderosa Dougl. ex Laws) ecosystem and in a moist Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) ecosystem. The fate of deuterated water applied to small plots to create a strong horizontal soil water potential gradient was also monitored to assess the potential for horizontal redistribution of water and utilization of redistributed water by co-occurring shallow-rooted plants. In a 20-year-old Douglas-fir stand, approximately 28% of the water removed daily from the upper 2 m of soil was replaced by nocturnal hydraulic redistribution during late August. In an old-growth ponderosa pine stand, approximately 35% of the total daily water utilization from the upper 2 m of soil appeared to be replaced by hydraulic redistribution during July and August. By late September, hydraulic redistribution in the ponderosa pine stand was no longer apparent, even though total water use from the upper 2 m of soil was nearly identical to that observed earlier. Based on these results, hydraulic redistribution would allow 21 and 16 additional days of stored water to remain in the upper soil horizons in the ponderosa pine and Douglas-fir stands, respectively, after a 60-day drought. At both sites, localized applications of deuterated water induced strong reversal of root sap flow and caused soil water content to cease declining or even temporarily increase at locations too distant from the site of water application to have been influenced by movement of water through the soil without facilitation by roots. Xylem water deuterium values of ponderosa pine seedlings suggested utilization of redistributed water. Therefore, hydraulic redistribution may enhance seedling survival and maintain overstory transpiration during summer drought. These first approximations of the extent of hydraulic redistribution in these ecosystems suggest that it is likely to be an important process in both wet and dry forests of the Pacific Northwest.     

Response of stomatal conductance to drought in ponderosa pine: implications for carbon and ozone uptake

To gain insight into the limitations imposed by a typical Mediterranean-climate summer drought on the uptake of carbon and ozone in the ponderosa pine (Pinus ponderosa Dougl. ex Laws.) ecosystem, we compared diurnal trends in leaf physiology of young trees in a watered and a control plot located in the Sierra Nevada Mountains, CA, USA (Blodgett Forest, 38 degrees 53' N, 120 degrees 37' W, 1315 m elevation). Predawn water potential of trees in the watered plot remained above -0.3 MPa throughout the growing season, whereas it dropped in the control plot from -0.24 to -0.52 MPa between late May and mid-August. Photosynthesis and stomatal conductance of trees in the watered plot were relatively insensitive to atmospheric vapor pressure deficit (VPD), whereas gas exchange of trees in the control plot varied with changes in soil water, VPD and temperature. Although the 1998 growing season was abnormally wet, we saw a pronounced drought effect at the control site. Over the 2 months following the onset of watering, carbon and ozone uptake were measured on three days at widely spaced intervals. Carbon uptake per unit leaf area by 1-year-old foliage of trees in the control plot was 39, 35 and 30% less, respectively, than in the watered plot, and estimated ozone deposition per unit leaf area (ozone concentration times stomatal conductance) was 36, 46 and 41% less.     

Differences in leaf gas exchange and water relations among species and tree sizes in an Arizona pine-oak forest

We compared leaf gas exchange and water potential among the dominant tree species and major size classes of trees in an upland, pine-oak forest in northern Arizona. The study included old-growth Gambel oak (Quercus gambelii Nutt.), and sapling, pole, and old-growth ponderosa pines (Pinus ponderosa var. scopulorum Dougl. ex Laws.). Old-growth oak had higher predawn leaf water potential (Psi(leaf)) than old-growth pine, indicating greater avoidance of soil water stress by oak. Old-growth oak had higher stomatal conductance (G(w)), net photosynthetic rate (P(n)), and leaf nitrogen concentration, and lower daytime Psi(leaf) than old-growth pine. Stomatal closure started at a daytime Psi(leaf) of about -1.9 MPa for pine, whereas old-growth oak showed no obvious reduction in G(w) at Psi(leaf) values greater than -2.5 MPa. In ponderosa pine, P(n) and G(w) were highly sensitive to seasonal and diurnal variations in vapor pressure deficit (VPD), with similar sensitivity for sapling, pole, and old-growth trees. In contrast, P(n) and G(w) were less sensitive to VPD in Gambel oak than in ponderosa pine, suggesting greater tolerance of oak to atmospheric water stress. Compared with sapling pine, old-growth pine had lower morning and afternoon P(n) and G(w), predawn Psi(leaf), daytime Psi(leaf), and soil-to-leaf hydraulic conductance (K(l)), and higher foliar nitrogen concentration. Pole pine values were intermediate between sapling and old-growth pine values for morning G(w) and daytime Psi(leaf), similar to sapling pine for predawn Psi(leaf), and similar to old-growth pine for morning and afternoon P(n), afternoon G(w), K(l), and foliar nitrogen concentration. For the pines, low predawn Psi(leaf), daytime Psi(leaf), and K(l) were associated with low P(n) and G(w). Our data suggest that hydraulic limitations are important in reducing P(n) in old-growth ponderosa pine in northern Arizona, and indicate greater avoidance of soil water stress and greater tolerance of atmospheric water stress by old-growth Gambel oak than by old-growth ponderosa pine.     

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.     

Stomata open at night in pole-sized and mature ponderosa pine: implications for O3 exposure metrics

Ponderosa pine (Pinus ponderosa Dougl. ex Laws.) is widely distributed in the western USA. We report the lack of stomatal closure at night in early summer for ponderosa pine at two of three sites investigated. Trees at a third site with lower nitrogen dioxide and nitric acid exposure, but greater drought stress, had slightly open stomata at night in early summer but closed stomata at night for the rest of the summer. The three sites had similar background ozone exposure during the summer of measurement (2001). Nighttime stomatal conductance (gs) ranged from one tenth to one fifth that of maximum daytime values. In general, pole-sized trees (< 40 years old) had greater nighttime gs than mature trees (> 250 years old). In late summer, nighttime gs was low (< 3.0 mmol H2O m(-2) s(-1)) for both tree size classes at all sites. Measurable nighttime gs has also been reported in other conifers, but the values we observed were higher. In June, nighttime ozone (O3) uptake accounted for 9, 5 and 3% of the total daily O3 uptake of pole-sized trees from west to east across the San Bernardino Mountains. In late summer, O3 uptake at night was < 2% of diel uptake at all sites. Nocturnal O3 uptake may contribute to greater oxidant injury development, especially in pole-sized trees in early summer.     

A comparison of three approaches to modeling leaf gas exchange in annually drought-stressed ponderosa pine forests

We tested, compared and modified three models of stomatal conductance at the leaf level in a forest ecosystem where drought stress is a major factor controlling forest productivity. The models were tested against 2 years (1999 and 2000) of leaf-level measurements on ponderosa pine (Pinus ponderosa Dougl. ex Laws.) growing in the Mediterranean climate of California, USA. The Ball, Woodrow and Berry (1987) (BWB) model was modified to account for soil water stress. Among the models, results of the modified BWB model were in the closest agreement with observations (r2 = 0.71). The Jarvis (1976) model showed systematic simulation errors related to vapor pressure deficit (r2 = 0.65). Results of the Williams, Rastetter, Fernandes et al. (1996) (SPA) model showed the poorest correlation with empirical data, but this model has only one calibration parameter (r2 = 0.60). Sensitivity analyses showed that, in all three models, predictions of stomatal conductance were most responsive to photosynthetically active radiation and soil water content. Stomatal conductance showed little sensitivity to vapor pressure deficit in the Jarvis model, whereas in both the BWB and SPA models, vapor pressure deficit (or relative humidity) was the third most important variable. Parameterization of the SPA model was in accordance with the parameterization of the modified BWB model, although the two models differ greatly. Measured and modeled results indicate that stomatal behavior is not water conservative during spring; however, during summer, when soil water content is low and vapor pressure deficit is high, stomatal conductance decreases and, according to the models, intrinsic water- use efficiency increases.     

Converging patterns of uptake and hydraulic redistribution of soil water in contrasting woody vegetation types

We used concurrent measurements of soil water content and soil water potential (Psi(soil)) to assess the effects of Psi(soil) on uptake and hydraulic redistribution (HR) of soil water by roots during seasonal drought cycles at six sites characterized by differences in the types and amounts of woody vegetation and in climate. The six sites included a semi-arid old-growth ponderosa pine (Pinus ponderosa Dougl. ex P. Laws & C. Laws) forest, a moist old-growth Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) forest, a 24-year-old Douglas-fir forest and three Brazilian savanna sites differing in tree density. At all of the sites, HR was confined largely to the upper 60 cm of soil. There was a common threshold relationship between the relative magnitude of HR and Psi(soil) among the six study sites. Below a threshold Psi(soil) of about -0.4 MPa, overnight recharge of soil water storage increased sharply, and reached a maximum value of 80-90% over a range of Psi(soil) from ~ -1.2 to -1.5 MPa. Although amounts of water hydraulically redistributed to the upper 60 cm of soil were relatively small (0 to 0.4 mm day(-1)), they greatly reduced the rates of seasonal decline in Psi(soil). The effectiveness of HR in delaying soil drying diminished with increasing sapwood area per ground area. The relationship between soil water utilization and Psi(soil) in the 20-60-cm layer was nearly identical for all six sites. Soil water utilization varied with a surrogate measure of rhizosphere conductance in a similar manner at all six sites. The similarities in relationships between Psi(soil) and HR, soil water utilization and relative rhizosphere conductance among the six sites, suggests that, despite probable differences in maximum rooting depth and density, there was a convergence in biophysical controls on soil water utilization and redistribution in the upper soil layers where the density of finer roots is greatest.     

Leaf area distribution and radiative transfer in open-canopy forests: implications for mass and energy exchange

Leaf area and its spatial distribution are key canopy parameters needed to model the radiation regime within a forest and to compute the mass and energy exchange between a forest and the atmosphere. A much larger proportion of available net radiation is received at the forest floor in open-canopy forests than in closed-canopy forests. The proportion of ecosystem water vapor exchange (lambda E) and sensible heat exchange from the forest floor is therefore expected to be larger in open-canopy forests than in closed-canopy forests. We used a combination of optical and canopy geometry measurements, and robust one- and three-dimensional models to evaluate the influence of canopy architecture and radiative transfer on estimates of carbon, water and energy exchange of a ponderosa pine (Pinus ponderosa Dougl. ex Laws.) forest. Three-dimensional model simulations showed that the average probability of diffuse and direct radiation transmittance to the forest floor was greater than if a random distribution of foliage had been assumed. Direct and diffuse radiation transmittance to the forest floor was 28 and 39%, respectively, in the three-dimensional model simulations versus 23 and 31%, respectively, in the one-dimensional model simulations. The assumption of randomly distributed foliage versus inclusion of clumping factors in a one-dimensional, multi-layer biosphere-atmosphere gas exchange model (CANVEG) had the greatest effect on simulated annual net ecosystem exchange (NEE) and soil evaporation. Assuming random distribution, NEE was 41% lower, net photosynthesis 3% lower, total lambda E 10% lower, and soil evaporation 40% lower. The same comparisons at LAI 5 showed a similar effect on annual NEE estimates (37%) and lambda E (12%), but a much larger effect on net photosynthesis (20%), suggesting that, at low LAI, canopies are mostly sunlit, so that redistribution of light has little effect on net photosynthesis, whereas the effect on net photosynthesis is much greater at high LAIs.     

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.     

Toward reference conditions: wildfire effects on flora in an old-growth ponderosa pine forest

Remote ponderosa pine (Pinus ponderosa) forests on the North Rim of Grand Canyon National Park, Arizona, USA provide valuable examples of reference conditions due to their relatively uninterrupted fire regimes, limited grazing history, and protection from logging. Wildfire is an important disturbance agent in upland forests of the Interior West, yet repeated measurements taken before and after lightning-ignited fires are rare. In 1999, a low-severity Wildland Fire Use fire burned 156 ha on Fire Point, a peninsula dominated by old-growth ponderosa pines, which had not burned for at least 76 years. We measured understory plant community and forest floor characteristics in 1998 (1 year before the fire) and 2001 (2 years after the fire) at this site and at nearby reference sites that did not burn in 1999 but have had continuing fire regimes throughout the past century. After the wildfire, the plant community at Fire Point shifted toward higher compositional similarity with the reference sites. Analysis of functional group composition indicated that this change was due primarily to an increase in annual and biennial forbs. Gayophytum diffusum, Polygonum douglasii, Chenopodium spp., Solidago spp., Elymus elymoides, Calochortus nuttallii, Hesperostipa comata, and Lotus spp. were indicative of forests influenced by recent fires. Species richness, plant cover, plant layer density and plant diversity were significantly lower at Fire Point than at the reference sites, possibly due to long-term fire exclusion, but the fire did not increase the rate of change in these variables after 2 years. Few exotic species were present at any site. Forest floor depths at Fire Point were reduced to depths similar to the reference sites, primarily due to consumption of the duff layer. There was a significant inverse relationship between the ratio of duff:litter and species richness. Compared to fire-excluded forests, old-growth ponderosa pine forests influenced by low-intensity surface fires generally have greater plant species richness (especially annual forbs) and lighter fuel loads. This study supports the continued application of the Wildland Fire Use strategy in old-growth montane forests to maintain and improve forest health by altering understory species composition and reducing fuel loads.     

Ecological lessons from long-term studies in experimental forests: Ponderosa pine silviculture at Pringle Falls Experimental Forest, central Oregon

The United States Department of Agriculture (USDA) Forest Service celebrated the 100th anniversary of its network of experimental forests in 2008. For a century, the network of experimental forests have contributed immensely—both in the US and around the world—to the practical understanding of the environment and to the formation of management approaches and policies that affect our use of forests and the natural resources they contain. Experimental forests provide places for long-term science and management studies in major vegetation types across the 789,140 km² of public land administered by the Forest Service. They also provide an incredible wealth of records and knowledge of environmental change in natural and managed forest ecosystems across the United States. Pringle Falls Experimental Forest (Pringle Falls), southwest of Bend, Oregon, is the oldest experimental forest in the Pacific Northwest and is the site of some of the earliest forest management and silviculture research in this region. Research at this site began in 1914, and it was formally established as part of the national network of experimental forests in 1931 as a center for silviculture, forest management, and insect and disease research in ponderosa pine forests east of the Oregon Cascade Range. As part of the celebration of the Centennial Anniversary of the network of experimental forests, it is timely to examine a few of the ecological lessons that have resulted from long-term studies at Pringle Falls. Two different yet interconnected themes are traced: (1) management of existing old-growth ponderosa pine; and (2) management of young or immature ponderosa pine. Examples of ecological lessons, drawn from long-term studies established or followed by James W. Barrett, Matt D. Busse, Patrick H. Cochran, Walter G. Dahms, F. Paul Keen, Ernest L. Kolbe, Edwin L. Mowat, William W. Oliver, Ram Oren, Chester T. Youngberg, and Jianwei Zhang illustrate how work at Pringle Falls has both pursued and influenced societal demands for forest management strategies, and how this trajectory has cycled back to the themes under which the experimental forest was first established. Finally, these two themes are integrated as drivers for new landscape-scale long-term research at Pringle Falls, designed to evaluate the effects of thinning and fuel reduction treatments on multiple, interacting forest stresses of fire, insects, wind, and climate change.     

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.     

Ozone uptake, water loss and carbon exchange dynamics in annually drought-stressed Pinus ponderosa forests: measured trends and parameters for uptake modeling

This paper describes 3 years of physiological measurements on ponderosa pine (Pinus ponderosa Dougl. ex Laws.) growing along an ozone concentration gradient in the Sierra Nevada, California, including variables necessary to parameterize, validate and modify photosynthesis and stomatal conductance algorithms used to estimate ozone uptake. At all sites, gas exchange was under tight stomatal control during the growing season. Stomatal conductance was strongly correlated with leaf water potential (R2=0.82), which decreased over the growing season with decreasing soil water content (R2=0.60). Ozone uptake, carbon uptake, and transpirational water loss closely followed the dynamics of stomatal conductance. Peak ozone and CO2 uptake occurred in early summer and declined progressively thereafter. As a result, periods of maximum ozone uptake did not correspond to periods of peak ozone concentration, underscoring the inappropriateness of using current metrics based on concentration (e.g., SUM0, W126 and AOT40) for assessing ozone exposure risk to plants in this climate region. Both Jmax (maximum CO2-saturated photosynthetic rate, limited by electron transport) and Vcmax (maximum rate of Rubisco-limited carboxylation) increased toward the middle of the growing season, then decreased in September. Intrinsic water-use efficiency rose with increasing drought stress, as expected. The ratio of Jmax to Vcmax was similar to literature values of 2.0. Nighttime respiration followed a Q10 of 2.0, but was significantly higher at the high-ozone site. Respiration rates decreased by the end of the summer as a result of decreased metabolic activity and carbon stores.     

Physiological responses of ponderosa pine in western Montana to thinning, prescribed fire and burning season

Low-elevation ponderosa pine (Pinus ponderosa Dougl. ex. Laws.) forests of the northern Rocky Mountains historically experienced frequent low-intensity fires that maintained open uneven-aged stands. A century of fire exclusion has contributed to denser ponderosa pine forests with greater competition for resources, higher tree stress and greater risk of insect attack and stand-destroying fire. Active management intended to restore a semblance of the more sustainable historic stand structure and composition includes selective thinning and prescribed fire. However, little is known about the relative effects of these management practices on the physiological performance of ponderosa pine. We measured soil water and nitrogen availability, physiological performance and wood radial increment of second growth ponderosa pine trees at the Lick Creek Experimental Site in the Bitterroot National Forest, Montana, 8 and 9 years after the application of four treatments: thinning only; thinning followed by prescribed fire in the spring; thinning followed by prescribed fire in the fall; and untreated controls. Volumetric soil water content and resin capsule ammonium did not differ among treatments. Resin capsule nitrate in the control treatment was similar to that in all other treatments, although burned treatments had lower nitrate relative to the thinned-only treatment. Trees of similar size and canopy condition in the three thinned treatments (with and without fire) displayed higher leaf-area-based photosynthetic rate, stomatal conductance and mid-morning leaf water potential in June and July, and higher wood radial increment relative to trees in control units. Specific leaf area, mass-based leaf nitrogen content and carbon isotope discrimination did not vary among treatments. Our results suggest that, despite minimal differences in soil resource availability, trees in managed units where basal area was reduced had improved gas exchange and growth compared with trees in unmanaged units. Prescribed fire (either in the spring or in the fall) in addition to thinning, had no measurable effect on the mid-term physiological performance and wood growth of second growth ponderosa pine.     

Tree mortality in drought-stressed mixed-conifer and ponderosa pine forests, Arizona, USA

We monitored tree mortality in northern Arizona (USA) mixed-conifer and ponderosa pine (Pinus ponderosa Dougl. ex Laws) forests from 1997 to 2007, a period of severe drought in this area. Mortality was pervasive, occurring on 100 and 98% of 53 mixed-conifer and 60 ponderosa pine plots (1-ha each), respectively. Most mortality was attributable to a suite of forest insects, mediated by drought stress. The number of trees dying from 2002 to 2007 was more than 200% greater than the number dying from 1997 to 2002 in mixed-conifer forest and 74% greater in ponderosa pine forest. Extent of mortality was spatially variable in both forest types. Median cumulative mortality (the ratio of dead to live trees) increased by approximately 53 and 65% in mixed-conifer and ponderosa pine forests, respectively, from 2002 to 2007. Median mortality rates from 2002 to 2007 were approximately 2.0% year⁻¹ in mixed-conifer forest (range = 0-28.5%) and 0.4% year⁻¹ in ponderosa pine forest (range = 0-13.6%). Mortality rates generally were not strongly related to either elevation or stand density. Mortality was nonrandom with respect to tree size classes and species. Proportions of trees dying were greatest in the largest size classes, particularly in mixed-conifer forest, where mortality in the largest size class exceeded 22% from 2002 to 2007. Mortality in mixed-conifer forest was particularly pronounced for quaking aspen (85%) and white fir (28%), the least drought tolerant species present. These results provide an early glimpse of how these forest types are likely to respond to predicted climate changes in the southwestern USA. They suggest that these forests are not resilient to climate change, and that treatments to increase resilience to climate change may be appropriate. Research on causes of spatial heterogeneity in extent of mortality might suggest valuable approaches to aid in increasing resilience.     

Perpetuating old ponderosa pine

We review current knowledge about the use of management treatments to reduce human-induced threats to old ponderosa pine (Pinus ponderosa) trees. We address the following questions: Are fire-induced damage and mortality greater in old than younger trees? Can management treatments ameliorate the detrimental effects of fire, competition-induced stress, and drought on old trees? Can management increase resistance of old trees to bark beetles? We offer the following recommendations for the use of thinning and burning treatments in old-growth ponderosa pine forests. Treatments should be focused on high-value stands where fire exclusion has increased fuels and competition and where detrimental effects of disturbance during harvesting can be minimized. Fuels should be reduced in the vicinity of old trees prior to prescribed burns to reduce fire intensity, as old trees are often more prone to dying after burning than younger trees. Raking the forest floor beneath old trees prior to burning may not only reduce damage from smoldering combustion under certain conditions but also increase fine-root mortality. Thinning of neighboring trees often increases water and carbon uptake of old trees within 1 year of treatment, and increases radial growth within several years to two decades after treatment. However, stimulation of growth of old trees by thinning can be negated by severe drought. Evidence from young trees suggests that management treatments that cause large increases in carbon allocation to radial xylem growth also increase carbon allocation to constitutive resin defenses against bark beetle attacks, but evidence for old trees is scarce. Prescribed, low-intensity burning may attract bark beetles and increase mortality of old trees from beetle attacks despite a stimulation of bole resin production.     

Belowground carbon pools and processes in different age stands of Douglas-fir

Forest floor material and soil organic matter may act as both a source and a sink in global CO2 cycles. Thus, the ecosystem processes controlling these pools are central to understanding the transfers of carbon (C) between the atmosphere and terrestrial systems. To examine these ecosystem processes, the effect of stand age on temporal carbon source-sink relationships was examined in 20-year-old, 40-year-old and old-growth stands of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) in the Cascade Mountains of south-central Washington State. Belowground C and nitrogen (N) storage and soil respiration were measured. In addition, nylon mesh bags containing homogenized soils from each site were buried at the respective sites to quantify root ingrowth and potential C sequestration and loss. The sites supporting the 20- and 40-year-old stands had soil C stores reflecting the C contributions from logging residue, coarse woody debris and stumps left after harvest. Because the N-fixer red alder (Alnus rubra Bong.) comprised 33% of the 40-year-old stand, this site had significantly greater concentrations and pools of N in the forest floor than sites without red alder. This N-rich site had consistently lower soil CO2 efflux rates during the growing season than the sites supporting the 20-year-old and old-growth stands. Estimated annual soil C efflux was 1367, 883 and 1194 g m-2 for the sites supporting the 20-, 40- and old-growth stands, respectively. These values are higher than previously reported values. Root ingrowth was significantly less in the 40-year-old stand than in the 20-year-old stand, and both young stands showed markedly less fine root growth than the old-growth stand. At the sites supporting the young stands, C and N were lost from the soil bags, whereas there was an increase in C and N in the soil bags at the site supporting the old-growth stand. The fine root growth and soil respiration data support the hypothesis that belowground C allocation decreases with increasing fertility. Quantification of the source-sink relationship of soil C at the three stands based on litterfall, relative root ingrowth and soil respiration measurements was compromised because of significant CO2 flux from decaying organic matter in the young stands.     

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.     

High temperature and drought stress effects on survival of Pinus ponderosa seedlings

We studied the effects of high temperature and drought on the survival, growth and water relations of seedlings of Pinus ponderosa (Dougl.) Lawson, one of few coniferous tree species that can successfully colonize drought-prone sites with high soil surface temperatures. Temperature profiles were measured with 0.07-mm thermocouples in a sparse ponderosa pine forest in northern Idaho. The soil surface and the adjacent 5 mm of air reached maximum temperatures exceeding 75 degrees C, well above the lethal temperature threshold for most plants. Air temperatures 50 mm above the soil surface (seedling needle height) rarely exceeded 45 degrees C. Pinus ponderosa seedlings that survived maintained basal stem temperatures as much as 15 degrees C lower than the surrounding air. The apparent threshold temperature at the seedling stem surface resulting in death was approximately 63 degrees C for less than 1 min. No correlation between seedling mortality and needle temperature was found, although some needles reached temperatures as high as 60 degrees C for periods of 相似文献   

Influence of ozone and nitrogen deposition on bark beetle activity under drought conditions

Four years of severe drought from 1999 through 2003 led to unprecedented bark beetle activity in ponderosa and Jeffrey pine in the San Bernardino and San Jacinto Mountains of southern California. Pines in the San Bernardino Mountains also were heavily impacted by ozone and nitrogenous pollutants originating from urban and agricultural areas in the Los Angeles basin. We studied bark beetle activity and bark beetle associated tree mortality in pines at two drought-impacted sites in the San Bernardino Mountains, one receiving high levels of atmospheric pollutants, and one with more moderate atmospheric input. We also investigated the effects of nitrogen addition treatments of 0, 50 and 150 kg N ha⁻¹ year⁻¹ at each site. Tree mortality and beetle activity were significantly higher at the high pollution site. Differences in beetle activity between sites were significantly associated with ozone injury to pines, while differences in tree mortality between sites were significantly associated with both ozone injury and fertilization level. Tree mortality was 9% higher and beetle activity 50% higher for unfertilized trees at the high pollution site compared to the low pollution site. Tree mortality increased 8% and beetle activity increased 20% under the highest rates of nitrogen additions at the low pollution site. The strong response in beetle activity to nitrogen additions at the low pollution site suggests that atmospheric nitrogen deposition increased tree susceptibility to beetle attack at the high deposition site. While drought conditions throughout the region were a major factor in decreased tree resistance, it appears that both ozone exposure and atmospheric nitrogen deposition further increased pine susceptibility to beetle attack.