Ozone exposure-response relationships parametrized for sixteen tree species with varying sensitivity in the United States.

It is well known that exposure to ambient O3 can decrease growth in many tree species in the United States (US). Our study reports experimental data from outdoor open-top chamber (OTC) studies that quantify total biomass response changes for seedlings of 16 species native to western and eastern North America, which were exposed to several levels of elevated O3 for one or more years. The primary objective of this study is to establish a reference set of parameters for these seedling exposure-response relationships using a 3-month (92 day) 12-hr W126 O3 metric used by US Environmental Protection Agency and other agencies to assess risk to trees from O3 exposure. We classified the 16 species according to their sensitivity, based on the biomass loss response functions to protect from a 5% biomass loss. The three-month 12-h W126 estimated to result in a 5% biomass loss was 2.5-9.2 ppm-h for sensitive species, 20.8-25.2 ppm-h for intermediate species, and > 28.7 ppm-h for insensitive species. The most sensitive tree species include black cherry, ponderosa pine, quaking aspen, red alder, American sycamore, tulip poplar and winged sumac. These species are ecologically important and widespread across US. The effects of O3 on whole-plant biomass depended on exposure duration and dynamics and on the number of successive years of exposure. These species-specific exposure-response relationships will allow US agencies and other groups to better estimate biomass losses based on ozone exposures in North America and can be used in risk assessment and scenario analyses.

Physiological responses of Douglas-fir to climate and forest disturbances as detected by cellulosic carbon and oxygen isotope ratios.

Swiss needle cast (SNC), caused by a fungal pathogen, Nothophaeocryptopus gaeumannii, is a major forest disease of Douglas-fir (Pseudotsuga menziesii) stands of the Pacific Northwest (PNW). There is mounting concern that the current SNC epidemic occurring in Oregon and Washington will continue to increase in severity, frequency and spatial extent with future warming. Nothophaeocryptopus gaeumannii occurs wherever its host is found, but very little is known about the history and spatial distribution of SNC and its effects on growth and physiological processes of mature and old-growth forests within the Douglas-fir region of the PNW. Our findings show that stem growth and physiological responses of infected Douglas-fir to climate and SNC were different between sites, growth periods and disease severity based on cellulosic stable carbon and oxygen isotope ratios and ring width data in tree rings. At a coastal Oregon site within the SNC impact zone, variations in stem growth and Δ13C were primarily influenced by disproportional reductions in stomatal conductance (gs) and assimilation (A) caused by a loss of functioning stomates through early needle abscission and stomatal occlusion by pseudothecia of N. gaeumannii. At the less severely infected inland sites on the west slopes of Oregon's Cascade Range, stem growth correlated negatively with δ18O and positively with Δ13C, indicating that gs decreased in response to high evaporative demand with a concomitant reduction in A. Current- and previous-years summer vapor pressure deficit was the principal seasonal climatic variable affecting radial stem growth and the dual stable isotope ratios at all sites. Our results indicate that rising temperatures since the mid-1970s has strongly affected Douglas-fir growth in the PNW directly by a physiological response to higher evaporative demand during the annual summer drought and indirectly by a major SNC epidemic that is expanding regionally to higher latitudes and higher elevations.

Regional patterns of increasing Swiss needle cast impacts on Douglas-fir growth with warming temperatures.

The fungal pathogen, Phaeocryptopus gaeumannii, causing Swiss needle cast (SNC) occurs wherever Douglas-fir is found but disease damage is believed to be limited in the U.S. Pacific Northwest (PNW) to the Coast Range of Oregon and Washington (Hansen et al., Plant Disease, 2000, 84, 773; Rosso & Hansen, Phytopathology, 2003, 93, 790; Shaw, et al., Journal of Forestry, 2011, 109, 109). However, knowledge remains limited on the history and spatial distribution of SNC impacts in the PNW. We reconstructed the history of SNC impacts on mature Douglas-fir trees based on tree-ring width chronologies from western Oregon. Our findings show that SNC impacts on growth occur wherever Douglas-fir is found and is not limited to the coastal fog zone. The spatiotemporal patterns of growth impact from SNC disease were synchronous across the region, displayed periodicities of 12-40 years, and strongly correlated with winter and summer temperatures and summer precipitation. The primary climatic factor limiting pathogen dynamics varied spatially by location, topography, and elevation. SNC impacts were least severe in the first half of the 20th century when climatic conditions during the warm phase of the Pacific Decadal Oscillation (1924-1945) were less conducive to pathogen development. At low- to mid-elevations, SNC impacts were most severe in 1984-1986 following several decades of warmer winters and cooler, wetter summers including a high summer precipitation anomaly in 1983. At high elevations on the west slope of the Cascade Range, SNC impacts peaked several years later and were the greatest in the 1990s, a period of warmer winter temperatures. Climate change is predicted to result in warmer winters and will likely continue to increase SNC severity at higher elevations, north along the coast from northern Oregon to British Columbia, and inland where low winter temperatures currently limit growth of the pathogen. Our findings indicate that SNC may become a significant forest health problem in areas of the PNW beyond the coastal fog zone.

Seasonal patterns of bole water content in old growth Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco).

Large conifer trees in the Pacific Northwest, USA (PNW) use stored water to extend photosynthesis, both diurnally and seasonally. This is particularly important during the summer drought, which is characteristic of the region. In the PNW, climate change is predicted to result in hotter, drier summers and warmer, wetter winters with decreased snowpack by mid-century. Understanding seasonal bole water dynamics in relation to climate factors will enhance our ability to determine the vulnerability of forests to climate change. Seasonal patterns of bole water content in old-growth Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) trees were studied in the Cascade Mountains of western Oregon, USA. Relative water content (RWC) was monitored hourly in three 400+ and three ~150 years-old trees using permanently mounted dielectric devices for 10 years. RWC increased during the late spring and early summer to maximum levels in August then decreased into fall and remained low over winter. The difference between minimum RWC in the winter and maximum in mid-summer averaged 4.5 and 2.3% for the older and younger trees, respectively, across all years. RWC closely followed growth and was positively correlated with air and soil temperature, vapor pressure deficit and photosynthetically active radiation, but lagged plant available soil water. The progressive decrease in RWC seen each year from mid-summer through fall was attributed to net daily loss of water during the summer drought. The marked increase in RWC observed from spring to mid-summer each year was hypothesized to be the period of embolism repair and water recharge in elastic tissues. We conclude that bole water content is an integral part of tree water dynamics enabling trees to extend carbon assimilation into drought periods and during periods when cold soil inhibits water uptake by roots, an adaptation that could benefit the survival of large PNW trees under climate change.

A likelihood-based time series modeling approach for application in dendrochronology to examine the growth-climate relations and forest disturbance history.

A time series intervention analysis (TSIA) of dendrochronological data to infer the tree growth-climate-disturbance relations and forest disturbance history is described. Maximum likelihood is used to estimate the parameters of a structural time series model with components for climate and forest disturbances (i.e., pests, diseases, fire). The statistical method is illustrated with a tree-ring width time series for a mature closed-canopy Douglas-fir stand on the west slopes of the Cascade Mountains of Oregon, USA that is impacted by Swiss needle cast disease caused by the foliar fungus, Phaecryptopus gaeumannii (Rhode) Petrak. The likelihood-based TSIA method is proposed for the field of dendrochronology to understand the interaction of temperature, water, and forest disturbances that are important in forest ecology and climate change studies.

Seasonal and long-term effects of CO2 and O3 on water loss in ponderosa pine and their interaction with climate and soil moisture.

Evapotranspiration (ET) is driven by evaporative demand, available solar energy and soil moisture (SM) as well as by plant physiological activity which may be substantially affected by elevated CO2 and O3. A multi-year study was conducted in outdoor sunlit-controlled environment mesocosm containing ponderosa pine seedlings growing in a reconstructed soil-litter system. The study used a 2 x 2 factorial design with two concentrations of CO2 (ambient and elevated), two levels of O3 (low and high) and three replicates of each treatment. The objective of this study was to assess the effects of chronic exposure to elevated CO2 and O3, alone and in combination, on daily ET. This study evaluated three hypotheses: (i) because elevated CO2 stimulates stomatal closure, O3 effects on ET will be less under elevated CO2 than under ambient CO2; (ii) elevated CO2 will ameliorate the long-term effects of O3 on ET; and (iii) because conductance (g) decreases with decreasing SM, the impacts of elevated CO2 and O3, alone and in combination, on water loss via g will be greater in early summer when SM is not limiting than to other times of the year. A mixed-model covariance analysis was used to adjust the daily ET for seasonality and the effects of SM and photosynthetically active radiation when testing for the effects of CO2 and O3 on ET via the vapor pressure deficit gradient. The empirical results indicated that the interactive stresses of elevated CO2 and O3 resulted in a lesser reduction in ET via reduced canopy conductance than the sum of the individual effects of each gas. CO2-induced reductions in ET were more pronounced when trees were physiologically most active. O3-induced reductions in ET under ambient CO2 were likely transpirational changes via reduced conductance because needle area and root biomass were not affected by exposures to elevated O3 in this study.

Elevated CO2 and O3 effects on fine-root survivorship in ponderosa pine mesocosms.

Atmospheric carbon dioxide (CO(2)) and ozone (O(3)) concentrations are rising, which may have opposing effects on tree C balance and allocation to fine roots. More information is needed on interactive CO(2) and O(3) effects on roots, particularly fine-root life span, a critical demographic parameter and determinant of soil C and N pools and cycling rates. We conducted a study in which ponderosa pine (Pinus ponderosa) seedlings were exposed to two levels of CO(2) and O(3) in sun-lit controlled-environment mesocosms for 3 years. Minirhizotrons were used to monitor individual fine roots in three soil horizons every 28 days. Proportional hazards regression was used to analyze effects of CO(2), O(3), diameter, depth, and season of root initiation on fine-root survivorship. More fine roots were produced in the elevated CO(2) treatment than in ambient CO(2). Elevated CO(2), increasing root diameter, and increasing root depth all significantly increased fine-root survivorship and median life span. Life span was slightly, but not significantly, lower in elevated O(3), and increased O(3) did not reduce the effect of elevated CO(2). Median life spans varied from 140 to 448 days depending on the season of root initiation. These results indicate the potential for elevated CO(2) to increase the number of fine roots and their residence time in the soil, which is also affected by root diameter, root depth, and phenology.

An approach for evaluating the effectiveness of various ozone air quality standards for protecting trees.

We demonstrate an approach for evaluating the level of protection attained using a variety of forms and levels of past, current, and proposed Air Quality Standards (AQSs). The U.S. Clean Air Act requires the establishment of ambient air quality standards to protect health and public welfare. However, determination of attainment of these standards is based on ambient pollutant concentrations rather than prevention of adverse effects. To determine if a given AQS protected against adverse effects on vegetation, hourly ozone concentrations were adjusted to create exposure levels that "just attain" a given standard. These exposures were used in combination with a physiologically-based tree growth model to account for the interactions of climate and ozone. In the evaluation, we used ozone concentrations from two 6-year time periods from the San Bernardino Mountains in California. There were clear differences in the level of vegetation protection achieved with the various AQSs. Based on modeled plant growth, the most effective standards were the California 8-hr average maximum of 70 ppb and a seasonal, cumulative, concentration-weighted index (SUM06), which if attained, resulted in annual growth reductions of 1% or less. Least effective was the 1-hr maximum of 120 ppb which resulted in a 7% annual reduction. We conclude that combining climate, exposure scenarios, and a process-based plant growth simulator was a useful approach for evaluating effectiveness of current or proposed air quality standards, or evaluating the form and/or level of a standard based on preventing adverse growth effects.

Elevated CO(2) and temperature alter net ecosystem C exchange in a young Douglas fir mesocosm experiment.

We investigated the effects of elevated CO(2) (EC) [ambient CO(2) (AC) + 190 ppm] and elevated temperature (ET) [ambient temperature (AT) + 3.6 degrees C] on net ecosystem exchange (NEE) of seedling Douglas fir (Pseudotsuga menziesii) mesocosms. As the study utilized seedlings in reconstructed soil-litter-plant systems, we anticipated greater C losses through ecosystem respiration (R(e)) than gains through gross photosynthesis (GPP), i.e. negative NEE. We hypothesized that: (1) EC would increase GPP more than R(e), resulting in NEE being less negative; and (2) ET would increase R(e) more than GPP, resulting in NEE being more negative. We also evaluated effects of CO(2) and temperature on light inhibition of dark respiration. Consistent with our hypothesis, NEE was a smaller C source in EC, not because EC increased photosynthesis but rather because of decreased respiration resulting in less C loss. Consistent with our hypothesis, NEE was more negative in ET because R(e) increased more than GPP. The light level that inhibited respiration varied seasonally with little difference among CO(2) and temperature treatments. In contrast, the degree of light inhibition of respiration was greater in AC than EC. In our system, respiration was the primary control on NEE, as EC and ET caused greater changes in respiration than photosynthesis.

Bole water content shows little seasonal variation in century-old Douglas-fir trees.

Purportedly, large Douglas-fir trees in the American Pacific Northwest use water stored in bole tissues to ameliorate the effects of seasonal summer drought, the water content of bole tissues being drawn down over the summer months and replenished during the winter. Continuous monitoring of bole relative water content (RWC) in two 110-120-year-old Douglas-fir trees with ThetaProbe impedance devices provided an integrated measure of phloem-sapwood water content over 4 years. Seasonal changes in RWC closely tracked cambial activity and wood formation, but lagged changes in soil water content by 2-3 months. The RWC in the combined phloem and sapwood markedly increased during earlywood production in the late spring and early summer to maximum values of 64-77% as plant available soil water (ASW) decreased by approximately 60%. With transition and latewood formation, RWC decreased to minimum values of 59-72%, even as ASW increased in the fall. The difference between minimum RWC in the fall and maximum RWC in midsummer was only approximately 5%. Seasonal changes in bole RWC corresponded to cambial phenology, although decreasing AWS appeared to trigger the shift from earlywood to latewood formation.

CO2 and N-fertilization effects on fine-root length, production, and mortality: a 4-year ponderosa pine study.

We conducted a 4-year study of juvenile Pinus ponderosa fine root (< or =2 mm) responses to atmospheric CO2 and N-fertilization. Seedlings were grown in open-top chambers at three CO2 levels (ambient, ambient+175 mumol/mol, ambient+350 mumol/mol) and three N-fertilization levels (0, 10, 20 g m(-2) year(-1)). Length and width of individual roots were measured from minirhizotron video images bimonthly over 4 years starting when the seedlings were 1.5 years old. Neither CO2 nor N-fertilization treatments affected the seasonal patterns of root production or mortality. Yearly values of fine-root length standing crop (m m(-2)), production (m m(-2) year(-1)), and mortality (m m(-2) year(-1)) were consistently higher in elevated CO2 treatments throughout the study, except for mortality in the first year; however, the only statistically significant CO2 effects were in the fine-root length standing crop (m m(-2)) in the second and third years, and production and mortality (m m(-2) year(-1)) in the third year. Higher mortality (m m(-2) year(-1)) in elevated CO2 was due to greater standing crop rather than shorter life span, as fine roots lived longer in elevated CO2. No significant N effects were noted for annual cumulative production, cumulative mortality, or mean standing crop. N availability did not significantly affect responses of fine-root standing crop, production, or mortality to elevated CO2. Multi-year studies at all life stages of trees are important to characterize belowground responses to factors such as atmospheric CO2 and N-fertilization. This study showed the potential for juvenile ponderosa pine to increase fine-root C pools and C fluxes through root mortality in response to elevated CO2.

Stricter ozone ambient air quality standard has beneficial effect on ponderosa pine in California.

Ambient air quality standards and control strategies are implemented to protect humans and vegetation from adverse effects. We used a process-based tree-growth model (TREGRO) to show that over the past 37 years, changes in O(3) exposure, with accompanying variation in climate, are reflected in changes in the growth of Pinus ponderosa Dougl. ex Laws. in the San Bernardino Mountains near Los Angeles, California, USA. Despite variation in temperature and precipitation over the study period (1963-1999), O(3) exposure consistently reduced simulated tree growth. Simulated growth reductions increased concurrent with increasing O(3) exposure. The maximum growth reduction occurred in 1979. As O(3) exposures decreased during the 1980s and 1990s, effects on growth also decreased. This implies that emission control strategies taken to reduce exposures to attain O(3) standards benefited P. ponderosa growth in the San Bernardino Mountains. This modeling approach provides a powerful tool for solving the difficult problem of evaluating regulatory effectiveness by simulating plant response using long-term climate and air pollution exposure records for a given region.

Monoterpene levels in needles of Douglas fir exposed to elevated CO2 and temperature.

Monoterpene levels in current year needles of Douglas fir (Pseudotsuga menziesii (Mirb.) Franco) seedlings were measured at the end of 4 years of exposure to ambient or elevated CO2 (+179 micro mol mol-1), and ambient or elevated temperature (+0.3.5;C). Eleven monoterpenes were identified and quantified using gas chromatography/flame ionization detector/mass spectroscopy, with eight of these compounds regularly occurring in all trees examined. Elevated CO2 exposure significantly reduced the levels for four of the eight main compounds in needles. Total monoterpene production was reduced by 52% (P < 0.05). Elevated temperature also reduced monoterpene levels (P < 0.07). The combination of elevated temperature and elevated CO2 resulted in a 64% reduction in total monoterpenes compared with needles on ambient temperature trees. Two-way anova showed no significant temperature-CO2 interaction. It is hypothesized that seasonal reductions in needle monoterpene pools under elevated CO2 and temperature conditions may be due to a combination of competing carbon sinks, including increased carbon flux through the roots.

Optimizing minirhizotron sample frequency for an evergreen and deciduous tree species.

• When using minirhizotrons to study fine dynamics in natural ecosystems, it is important to determine how sample collection frequency influences estimates of fine root production and mortality. Minirhizotron images were collected twice per week from mature Pseudotsuga menziesii and Tilia cordata trees and analyzed to estimate fine root production and mortality. These data were used to create data sets reflecting sample frequencies of 1, 2, 4 or 8 wk. • When the sampling interval is long, fine roots can appear and disappear between samplings, leading to underestimates of production and mortality. For example, with an 8-wk sample frequency, 24 and 35% of the fine root production in P. menziesii and T. cordata , respectively, is not measured. Fine root mortality displays the same sensitivity to sample frequency. • Our experimental observations supported the previously published simulation analysis, which provides an estimate of the proportion of fine roots missed at different sample frequencies and is a tool that can be used to select a sample frequency to balance production and mortality accuracy with sampling and analytical effort.

Effects of elevated CO(2) and temperature on cold hardiness and spring bud burst and growth in Douglas-fir (Pseudotsuga menziesii).

We examined effects of elevated CO(2) and temperature on cold hardiness and bud burst of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) seedlings. Two-year-old seedlings were grown for 2.5 years in semi-closed, sunlit chambers at either ambient or elevated (ambient + ~ 4 degrees C) air temperature in the presence of an ambient or elevated (ambient + ~ 200 ppm) CO(2) concentration. The elevated temperature treatment delayed needle cold hardening in the autumn and slowed dehardening in the spring. At maximum hardiness, trees in the elevated temperature treatment were less hardy by about 7 degrees C than trees in the ambient temperature treatment. In general, trees exposed to elevated CO(2) were slightly less hardy during hardening and dehardening than trees exposed to ambient CO(2). For trees in the elevated temperature treatments, date to 30% burst of branch terminal buds was advanced by about 6 and 15 days in the presence of elevated CO(2) and ambient CO(2), respectively. After bud burst started, however, the rate of increase in % bud burst was slower in the elevated temperature treatments than in the ambient temperature treatments. Time of bud burst was more synchronous and bud burst was completed within a shorter period in trees at ambient temperature (with and without elevated CO(2)) than in trees at elevated temperature. Exposure to elevated temperature reduced final % bud burst of both leader and branch terminal buds and reduced growth of the leader shoot. We conclude that climatic warming will influence the physiological processes of dormancy and cold hardiness development in Douglas-fir growing in the relatively mild temperate region of western Oregon, reducing bud burst and shoot growth.

Effects of elevated CO(2) and nitrogen on the synchrony of shoot and root growth in ponderosa pine.

We monitored effects of elevated CO(2) and N fertilization on shoot and fine root growth of Pinus ponderosa Dougl. ex P. Laws. and C. Laws. grown in native soil in open-top field-exposure chambers at Placerville, CA, over a 2-year period. The experimental design was a replicated 3 x 3 factorial with the center treatment missing; plants were exposed to ambient (~365 micro mol mol(-1)) air or ambient air plus either 175 or 350 micro mol mol(-1) CO(2) in combination with one of three rates of N addition (0, 100 or 200 kg ha(-1) year(-1)). All CO(2) by N interactions were nonsignificant. Both the CO(2) and N treatments increased plant height, stem diameter and leaf area index (LAI). Elevated CO(2) increased fine root area density and the occurrence of mycorrhizae, whereas N fertilization increased coarse root area density but had no effect on fine root area density. Spring flushes of shoot height and diameter growth were initiated concurrently with the increase in new root area density but height and diameter growth reached their maxima before that of fine roots. The temporal patterns of root and shoot growth were not altered by providing additional CO(2) or N. Greatest root loss occurred in the summer, immediately following the period of greatest new fine root growth. Elevated N initially reduced the fine root area density/LAI ratio independently of CO(2) treatment, indicating that the relationship between fine roots and needles was not changed by CO(2) exposure.