Air Pollution Impacts to Some Important Species of Pine

In the past, many species of pine have been severely damaged by acute forms of air pollution associated with ore smelting. More recently, damage to ponderosa pine in the state of Washington has been associated with atmospheric fluoride, and this species in particular has also suffered smog damage in southern California. In the East, white pine has been declining in the vicinity of certain soft-coal-burning power plants, and in one case downwind from an oil-burning power plant: also, throughout the East, an atmospheric oxidant of unknown source has caused a type of needle blight on white pine. Trash burners and city dump combustion have led to the hilling of certain pine species, leaving other intermixed pine species unharmed. Localized damage to white pine from automobile exhaust has been observed. Differences in the susceptibility of neighboring trees of the same species to a given air pollutant have been observed repeatedly.     

Isozyme markers associated with O(3) tolerance indicate shift in genetic structure of ponderosa and Jeffrey pine in Sequoia National Park, California

Effects of canopy ozone (O(3)) exposure and signatures of genetic structure using isozyme markers associated with O(3) tolerance were analyzed in approximately 20-, approximately 80-, and >200-yr-old ponderosa (Pinus ponderosa Dougl. ex Laws.) and Jeffrey pine (Pinus jeffreyi Grev. & Balf.) in Sequoia National Park, California. For both species, the number of alleles and genotypes per loci was higher in parental trees relative to saplings. In ponderosa pine, the heterozygosity value increased, and the fixation index indicated reduction of homozygosity with increasing tree age class. The opposite tendencies were observed for Jeffrey pine. Utilizing canopy attributes known to be responsive to O(3) exposure, ponderosa pine was more symptomatic than Jeffrey pine, and saplings were more symptomatic than old growth trees. We suggest that these trends are related to differing sensitivity of the two species to O(3) exposure, and to higher O(3) exposures and drought stress that younger trees may have experienced during germination and establishment.     

Ozone Dosage Response of Ponderosa Pine Seedlings

Photochemical oxidant injury to ponderosa pine (Pinus ponderosa Laws) is a severe problem in the southern California mountains. Three-year-old ponderosa pines fumigated in controlled environment chambers with ozone at 0.15, 0.30, or 0.45 ppm had apparent photosynthesis rates reduced by 10, 70, and 85%, respectively, after 30 days exposure. A fumigation with 0.30 ppm ozone for 33 days depressed the cold perchloric acid extracted polysaccharides of both current and one-year-old needles by 40%. The 80% ethanol soluble sugar concentration of current year, ozone-injured needles increased 16% and that of the one-year-old needles decreased slightly. Both carbohydrate fractions of control trees in carbon-filtered air increased moderately. Higher, endogenous concentrations of ascorbic acid in needles did not protect the tissue from ozone injury. Apparent photosynthesis rate was a sensitive index for ozone dosage response. Needle carbohydrate depletion probably induces premature abscission.     

Effects of acidic fog on seedlings of Pinus ponderosa and Abies concolor: foliar injury, physiological and biochemical responses

Seedlings of Pinus ponderosa (ponderosa pine) and Abies concolor (white fir) were exposed to acidic fog (pH 2.0, 3.0 or 4.0) in open-field plots for six weeks. The two species exhibited dissimilar injury responses; neither current year nor previous year needles of ponderosa pine were injured by pH 2.0 fog, but current year needles exhibited higher membrane permeability responses (i.e. needle extract conductivity, K+ concentration). In comparison, both needle age classes in white fir were significantly injured by pH 2.0 fog, but no significant effects on membrane permeability were observed. For both species, whole-study average rates of net photosynthesis in previous year needles were lower in plants exposed to pH 2.0 fog than in plants treated with pH 4.0 fog. While decreased process rates coincided with leaf necrosis in white fir, stomatal closure appeared to be the mechanism of inhibition in ponderosa pine with pH 2.0 fog (i.e. no visible injury). The findings of the present study provide evidence that frequent applications of highly acidic fog (i.e. pH 2.0-3.0) can cause temporal alterations in membrane permeability and gas exchange rates in western conifer seedlings, in the presence or absence of visible injury. However, because incipient effects on other measures of foliage health were species-specific (i.e. concentrations of starch, photosynthetic pigments, inorganic nutrients), a general mechanism of phytotoxicity could not be identified.     

An Evaluation of the Effects of Ozone Injury on Radial Growth of Ponderosa Pine (Pinus ponderosa) in the Southern Sierra Nevada

Growth of ponderosa pines with visible symptoms of ozone injury was compared with that of asymptomatic trees in the southern Sierra Nevada, California. Time series analysis indicated that there was no significant reduction in annual radial increment of symptomatic trees during recent years compared to past growth and growth of asymptomatic trees. First order autocorrelation and climatic variables accounted for a large proportion of the variance in growth index, and winter precipitation was positively correlated with growth for all size and age classes. Although ozone concentrations are high enough to cause chlorosis and premature needle senescence in ponderosa pine, there has been no significant change in growth associated with ozone injury.     

Nitrate reductase activity as an indicator of ponderosa pine response to atmospheric nitrogen deposition in the San Bernardino Mountains

Determinations of nitrate reductase (NR) activity in ponderosa pine (Pinus ponderosa Dougl. ex. Laws.) needles were performed during summer 1994 in two areas (consisting of six different sites) with different nitrogen (N) deposition levels in the San Bernardino Mountains, southern California. Nitrate reductase activity was used as an integrative indicator of atmospheric nitrogen deposition to pine trees (direct uptake of N species from the atmosphere and N transported from the soil). Deposition of nitrate (NO3-) to pine branches was measured in order to determine dry atmospheric inputs of the oxidized N species to tree foliage. High NR activity was detected in all of the experimental sites. Activity of the enzyme was significantly higher at the locations characterized by higher NO3- deposition to branches--slight positive correlation between branch deposited NO3- and NR activity was found. However, high variability of NR in time and between the experimental sites discredit the NR assay as a reliable indicator of N deposition for ponderosa pine in the field conditions. This could be caused by substantial interference from other abiotic and biotic factors with tropospheric ozone as probably the most important one.     

Dry deposition of nitrogen and sulfur to Ponderosa and Jeffrey pine in the San Bernardino national forest in Southern California

Little is known about the concentrations, deposition rates, and effects of nitrogenous and sulfurous compounds in photochemical smog in the San Bernardino National Forest (SBNF) in southern California. Dry deposition of NO(3)(-) and NH(4)(+) to foliage of ponderosa pine (Pinus ponderosa Laws.) and Jeffrey pine (Pinus jeffreyi Grev. & Balf.) was correlated (R = 0.83-0.88) with historical average hourly O(3) concentations at 10 sites across an O(3) gradient in the SBNF. Mean deposition fluxes of NO(3)(-) to ponderosa and Jeffrey pine branches were 0.82 nmol M(-2)s(-1) at Camp Paivika (CP), a high-pollution site, and 0.19 nmol m(-2) s(-1) at Camp Osceola (CAO), a low-pollution site. Deposition fluxes of NH(4)(+) were 0.32 nmol m(-2) s(-1) at CP and 0.17 nmol m(-2) s(-1) at CAO, while mean values for SO(4)(2-) were 0.03 at CP and 0.02 nmol m(-2) s(-1) at CAO. Deposition fluxes to paper and nylon filters were higher in most cases than fluxes to pine branches at the same site. The results of this study suggest that an atmospheric concentration and deposition gradient of N and S compounds occurs along with the west-east O(3) gradient in the SBNF. Annual stand-level dry deposition rates for S and N at CP and CAO were estimated. Further studies are needed to determine if high N deposition loads in the SBNF significantly affect plant/soil nutrient relations, tree health, and the response of ponderosa pine to ozone.     

Interactive effects of CO2 and O3 on a ponderosa pine plant/litter/soil mesocosm.

To study individual and combined impacts of two important atmospheric trace gases, CO2 and O3, on C and N cycling in forest ecosystems; a multi-year experiment using a small-scale ponderosa pine (Pinus ponderosa Laws.) seedling/soil/litter system was initiated in April 1998. The experiment was conducted in outdoor, sun-lit chambers where aboveground and belowground ecological processes could be studied in detail. This paper describes the approach and methodology used, and presents preliminary data for the first two growing seasons. CO2 treatments were ambient and elevated (ambient + 280 ppm). O3 treatments were elevated (hourly averages to 159 ppb, cumulative exposure > 60 ppb O3, SUM 06 approximately 10.37 ppm h), and a low control level (nearly all hourly averages <40 ppb. SUM 06 approximately 0.07 ppm h). Significant (P < 0.05) individual and interactive effects occurred with elevated CO2 and elevated O3. Elevated CO2 increased needle-level net photosynthetic rates over both seasons. Following the first season, the highest photosynthetic rates were for trees which had previously received elevated O3 in addition to elevated CO2. Elevated CO2 increased seedling stem diameters, with the greatest increase at low O3. Elevated CO2 decreased current year needle % N in the summer. For 1-year-old needles measured in the fall there was a decrease in % N with elevated CO2 at low O3, but an increase in % N with elevated CO2 at elevated O3. Nitrogen fixation (measured by acetylene reduction) was low in ponderosa pine litter and there were no significant CO2 or O3 effects. Neither elevated CO2 nor elevated O3 affected standing root biomass or root length density. Elevated O3 decreased the % N in coarse-fine (1-2 mm diameter) but not in fine (< 1 mm diameter) roots. Both elevated CO2 and elevated O3 tended to increase the number of fungal colony forming units (CFUs) in the AC soil horizon, and elevated O3 tended to decrease bacterial CFUs in the C soil horizon. Thus, after two growing seasons we showed interactive effects of O3 and CO2 in combination, in addition to responses to CO2 or O3 alone for a ponderosa pine plant/litter/soil system.     

Empirical and simulated critical loads for nitrogen deposition in California mixed conifer forests

Empirical critical loads (CL) for N deposition were determined from changes in epiphytic lichen communities, elevated NO(3)(-) leaching in streamwater, and reduced fine root biomass in ponderosa pine (Pinus ponderosa Dougl. ex Laws.) at sites with varying N deposition. The CL for lichen community impacts of 3.1kg ha(-1) year(-1) is expected to protect all components of the forest ecosystem from the adverse effects of N deposition. Much of the western Sierra Nevada is above the lichen-based CL, showing significant changes in lichen indicator groups. The empirical N deposition threshold and that simulated by the DayCent model for enhanced NO(3)(-)leaching were 17kg N ha(-1) year(-1). DayCent estimated that elevated NO(3)(-) leaching in the San Bernardino Mountains began in the late 1950s. Critical values for litter C:N (34.1), ponderosa pine foliar N (1.1%), and N concentrations (1.0%) in the lichen Letharia vulpina ((L.) Hue) are indicative of CL exceedance.     

Variation in CO2 assimilation rate induced by simulated dew waters with different sources of hydroxyl radical (*OH) on the needle surfaces of Japanese red pine (Pinus densifora Sieb. et Zucc.)

The hydroxyl radical (*OH) is generated in polluted dew on the needle surfaces of Japanese red pine (Pinus densiflora Sieb. et Zucc.). This free radical, which is a potent oxidant, is assumed to be a cause of ecophysiological disorders of declining trees on the urban-facing side of Mt. Gokurakuji, western Japan. Mists of *OH-generating N(III) (HNO2 and NO2-) and HOOH + Fe + oxalate solutions (50 and 100 microM, pH 5.1-5.4) simulating the dew water were applied to the foliage of pine seedlings grown in open-top chambers in the early morning. Needles treated with 100 microM N(III) tended to have a greater maximum CO2 assimilation rate (Amax), a greater stomatal conductance (g(s)) and a greater needle nitrogen content (Nneedle), suggesting that N(III) mist acts as a fertilizer rather than as a phytotoxin. On the other hand, needles treated with 100 microM HOOH + Fe + oxalate solution showed the smallest Amax, g(s), and Nneedle, suggesting that the combination of HOOH + Fe + oxalate caused a decrease in needle productivity. The effects of HOOH + Fe + oxalate mist on pine needles were very similar to the symptoms of declining trees at Mt. Gokurakuji.     

Air Pollution Damage to the Forests of the Sierra Nevada Mountains of California

Numerous oxidant-injured ponderosa pine and other tree species have been detected in the lower elevation forests of the southern Sierra Nevada mountain range of California in the Sequoia National Forest, Sequoia-Kings Canyon National Parks, and on state and private lands. Affected pines were observed in a broad band following the many ridges that rise above the Central Valley floor to the 1900-2600 m elevation. Slight symptoms of ozone chlorotic mottle on ponderosa pine appeared from about 1000 m altitude, increased in severity as elevation increased and then diminished in severity above 2600 m. An apparent east to west gradient of disease was discovered; severity lessened as distance from the Central Valley increased. Atmospheric monitoring in three forested sites suggests that ozone occurs in doses sufficient to elicit symptoms of this widespread disease.     

Glutathione status and glutathione reductase activity in spruce needles of healthy and damaged trees at two mountain sites

Levels of glutathione, in both reduced and oxidized form, and glutathione reductase activity were monitored in needles of healthy and damaged spruce trees (Picea abies (L.) Karst.) during the course of four vegetation periods at two natural sites. The glutathione content and glutathione reductase activity showed a pronounced annual rhythm in undamaged trees, whereas damaged spruce trees deviated significantly from this course. In comparison with undamaged trees, damaged trees showed markedly increased levels of glutathione during the test period of 1989-1991. However, glutathione reductase activity differed in damaged and undamaged trees, only in 1989-1990. The ratio of reduced to oxidized glutathione (GSH/GSSG ratio) was slightly higher in damaged trees, and the highest levels were found during the winter months. In the case of damaged trees, a correlation between GSH/GSSG ratio and current ozone levels at the sites could be clearly established. The present results indicate that damaged trees suffer from increased oxidative stress, especially in the period from June to October.     

Trace element concentration in tree-rings biomonitoring centuries of environmental change

Inductively coupled plasma mass spectrometry (ICP-MS) was used to examine trace element concentration in tree-rings over three and half centuries to assess macro-trends of environmental change. Tree-rings of a 350+ year old mammoth ponderosa pine (Pinus ponderosa) were analyzed for element concentration and evaluated versus local and global historical events. The ponderosa pine was located 100 miles south of the Canada/USA border and 180 miles east of the Pacific Ocean, and grew near apple orchards, a public road, and Swakane Creek in western Washington, USA. The elements tested did not all display the same time versus concentration patterns. Copper and chromium displayed cyclic concentration patterns over the last 350+ years, which appear to be associated with local events. Strontium, barium, zinc and cadmium were found to be relatively constant between the mid 1600s and the early 1800s. Strontium, barium, zinc, and cadmium then increased beginning in the early 1800s for approximately 50 years then decreased to present day 2000. Significantly, similar changes seen in Ca, Mg, and Zn in other studies have been attributed to acid rain, whereas, in our study area there is no history of anthropogenic acid rain. Most importantly, our data goes back to the mid-1600s several hundred years further back than most other studies of this nature. This additional time data provides for a better context of trend data not previously available.     

Scots pine (Pinus sylvestris L.) growth and condition in a polluted environment: from decline to recovery

The results of long-term investigations of Scots pine (Pinus sylvestris L.) growth and condition in the impact zone of one of the biggest air pollution sources in Lithuania--mineral fertilizers plant "Achema" are presented. The main attention is laid to the recovery of damaged stands since annual emissions to air were reduced essentially. The investigations indicated, that the recovery of tree increment was mostly caused by distinct reduction of emissions of nitrogen and sulphur oxides and dust of mineral fertilizers. Despite reduced pollution, crown defoliation of investigated stands has continued to increase for a certain period. After the crown recovery of damaged stands has started, the recovery of most damaged survived trees was most intensive and convergence of defoliated to a different extend stands and trees is characteristic feature of this period. No defoliation threshold has been determined beyond of which recovery of trees would be impossible. Recovery of more than a quarter of damaged trees was registered even in the case of 90% of defoliation. Recovery of dominant trees occurs to be faster of that for suppressed trees within the same level of defoliation. The impact of stand density on the crown recovery rate is negative, the higher density (more intensive competition), the slower recovery of damaged trees. The dependence of growth rate on defoliation was found to be of logistic character: while crown defoliation consists up to 25-30%, tree increment losses are rather inconsiderable, further increase of defoliation leads to the essentially higher increment losses, however having achieved 65-70% defoliation, further increase of defoliation does not result such intensive decrease of radial increment.     

Contents of free amino acids in needles of Norway Spruce trees in relation to novel forest decline. Studies on trees from a site in the northern Black Forest

The free amino acid content in needles of Norway spruce trees (about 45-year-old) was determined by means of HPLC. The studied trees have been growing at a forestry site in the Black Forest which is characterized by a high impact of ozone and magnesium deficiency. Measurements were carried out on visibly healthy and on damaged trees on several dates during two vegetation periods and during the course of a day. The amino acids occurring at the highest concentrations were glutamate, arginine, aspartate, and glutamine. The typical seasonal changes in the content of free amino acids, with their minimum during summer, were disturbed in the needles of the damaged trees. Particularly in summer and autumn the damaged trees showed increased levels of total amino acids and of most single amino acids. During the course of a day, the needles of the damaged trees showed a reduced amplitude in diurnal variations with the absolute level almost always higher in comparison to that of the undamaged trees. The results suggest that a shift in protein turnover towards enhanced degradation has taken place.     

Selection effects of air pollution on gene pools of Norway spruce, European silver fir and European beech.

The effects of industrial pollution on allelic and genotypic structures of Norway spruce. European silver fir and European beech were investigated by means of isozyme analysis. In a mixed Norway spruce-silver fir forest stand in an area heavily polluted by sulphur dioxide and heavy metals in the region of Spis (eastern Slovakia), pairs of neighbouring damaged and apparently healthy trees were selected in two replicates (44 and 69 pairs in a heavily and moderately damaged stand, respectively). Pairwise sampling of trees with contrasting vitality was applied to reduce potential effects of site heterogeneity on the vitality of sampled trees. No significant differences in allelic and genotypic frequencies were found between sets of healthy and declining trees. There were differences in the single-locus heterozygosities, but these were not consistent between the replicates. However, the set of damaged trees exhibited higher levels of genetic multiplicity and diversity, possibly due to the deleterious effect of rare alleles under the conditions of air pollution. Consequently. following the decline of pollutant-sensitive trees, the remaining stand will be depleted of a part of alleles with unknown adaptive value to future selection pressures.     

Tree decline in North America

Air-borne, phytotoxic pollutants are known to adversely affect forest tree growth in North America. On a local scale, exposure to high concentrations of toxic gases, such as sulphur dioxide and fluoride, result in foliar injury, branch dieback, reduced radial growth and increased mortality in a variety of tree species. On a regional scale, the photochemical oxidant, ozone, is responsible for growth decline of pollutant-sensitive eastern white pine genotypes in the eastern United States, and of ponderosa and Jeffrey pines in the mountains of southern and central California. The etiology associated with the reported declines of high elevation red spruce in the Appalachian Mountains and of sugar maple in the northeastern United States and southeastern Canada is incompletely known at present. A complex of predisposing and inciting factors, including temperature and moisture stress, edaphic conditions, aluminium toxicity, insect depredation, and air-borne pollutants are probably involved in these declines.     

An evaluation of ozone exposure metrics for a seasonally drought-stressed ponderosa pine ecosystem

Ozone stress has become an increasingly significant factor in cases of forest decline reported throughout the world. Current metrics to estimate ozone exposure for forest trees are derived from atmospheric concentrations and assume that the forest is physiologically active at all times of the growing season. This may be inaccurate in regions with a Mediterranean climate, such as California and the Pacific Northwest, where peak physiological activity occurs early in the season to take advantage of high soil moisture and does not correspond to peak ozone concentrations. It may also misrepresent ecosystems experiencing non-average climate conditions such as drought years. We compared direct measurements of ozone flux into a ponderosa pine canopy with a suite of the most common ozone exposure metrics to determine which best correlated with actual ozone uptake by the forest. Of the metrics we assessed, SUM0 (the sum of all daytime ozone concentrations > 0) best corresponded to ozone uptake by ponderosa pine, however the correlation was only strong at times when the stomata were unconstrained by site moisture conditions. In the early growing season (May and June). SUM0 was an adequate metric for forest ozone exposure. Later in the season, when stomatal conductance was limited by drought. SUM0 overestimated ozone uptake. A better metric for seasonally drought-stressed forests would be one that incorporates forest physiological activity, either through mechanistic modeling, by weighting ozone concentrations by stomatal conductance, or by weighting concentrations by site moisture conditions.     

Occurrence of spruce bark beetles in forest stands at different levels of air pollution stress

The spruce bark beetle, Ips typographus (L.) is the most serious pest of mature spruce stands, mainly Norway spruce, Picea abies (L.) Karst. throughout Eurasia. A complex of weather-related events and other environmental stresses are reported to predispose spruce stands to bark beetle attack and subsequent tree mortality; however the possible role of industrial pollution as a predisposing factor to attack by this species is poorly understood. The abundance and dynamics of I. typographus populations was evaluated in 60-80 year old Norway spruce stands occurring on 10 x 50 ha sites in five countries within the Carpathian range that were selected in proximity to established ozone measurement sites. Data were recorded on several parameters including the volume of infested trees, captures of adult beetles in pheromone traps, number of attacks, and the presence and relative abundance of associated bark beetle species. In several cases, stands adjacent to sites with higher ozone values were associated with higher bark beetle populations. The volume of sanitary cuttings, a reflection of tree mortality, and the mean daily capture of beetles in pheromone traps were significantly higher at sites where the O(3) level was higher. However, the mean infestation density on trees was higher in plots associated with lower O(3) levels. Captures of beetles in pheromone traps and infestation densities were higher in the zone above 800 m. However, none of the relationships was conclusive, suggesting that spruce bark beetle dynamics are driven by a complex interaction of biotic and abiotic factors and not by a single parameter such as air pollution.     

Growth and photosynthetic responses of two pine species (Pinus koraiensis and Pinus rigida) in a polluted industrial region in Korea

We investigated the effects of pollutants on two pine species (Pinus koraiensis and Pinus rigida) in an industrial region in Korea, using a physiological approach. The concentrations of fluorine (F) and chlorine (Cl) in the atmosphere, in precipitation and soil water at the damaged site were all significantly higher than at a control site. Moreover, the concentrations of F, Cl and Mn in pine needles were significantly higher, and essential elements and chlorophyll in needles were significantly lower at the damaged site than at the control site. The photosynthetic capacities, shoot length and survival statistics of needles of the two pines were all significantly reduced at the damaged site compared to the control site, especially P. rigida. Based on our comparison of photosynthetic responses and the concentrations of F, Cl and Mn in needles of the two pine species, P. koraiensis is more resistant to excess Mn in its needles than P. rigida.