Annual and spatial variation in shoot demography associated with masting in Betula grossa: comparison between mature trees and saplings

Backgrounds and Aims

Shoot demography affects the growth of the tree crown and the number of leaves on a tree. Masting may cause inter-annual and spatial variation in shoot demography of mature trees, which may in turn affect the resource budget of the tree. The aim of this study was to evaluate the effect of masting on the temporal and spatial variations in shoot demography of mature Betula grossa.

Methods

The shoot demography was analysed in the upper and lower parts of the tree crown in mature trees and saplings over 7 years. Mature trees and saplings were compared to differentiate the effect of masting from the effect of exogenous environment on shoot demography. The fate of different shoot types (reproductive, vegetative, short, long), shoot length and leaf area were investigated by monitoring and by retrospective survey using morphological markers on branches. The effects of year and branch position on demographic parameters were evaluated.

Key Results

Shoot increase rate, production of long shoots, bud mortality, length of long shoots and leaf area of a branch fluctuated periodically from year to year in mature trees over 7 years, in which two masting events occurred. Branches within a crown showed synchronized annual variation, and the extent of fluctuation was larger in the upper branches than the lower branches. Vegetative shoots varied in their bud differentiation each year and contributed to the dynamic shoot demography as much as did reproductive shoots, suggesting physiological integration in shoot demography through hormonal regulation and resource allocation.

Conclusions

Masting caused periodic annual variation in shoot demography of the mature trees and the effect was spatially variable within a tree crown. Since masting is a common phenomenon among tree species, annual variation in shoot demography and leaf area should be incorporated into resource allocation models of mature masting trees.     

Intra- and Extrabud Stages of Elementary Shoot Development in Ligneous Plants

Length of elementary shoots and the number of metamers in them have been compared in a number of ligneous plants. The shoot length varies to a greater extent than the number of metamers. The number of metamers with bud scales is the most invariable and species-specific. This indicates relative independence of the intrabud stage of the shoot development.     

两种亚热带榕属乔木植冠损伤后的位置依赖性枝发生

三峡库区需要大量植株用于三峡工程所涉及的诸多建设(如铁路、公路、建筑)之后的植被恢复和绿化.由于具有优美树冠和耐瘠薄土壤的能力,榕(Ficus microcarpa L.)和黄桷树(Fvirens Ait.var.sublanceolata(Miq.)Cornor) 在三峡库区作为绿化和行道树木大量栽植.在三峡库区,这两种树种苗木的培育主要通过切枝扦插的方式进行.大量切枝损伤植株植冠并且使叶组织数量减少,对植株生长有很大影响.植株生长与植株的枝发生格局有很大关系,为明确植冠损伤对植株生长的可能影响,对榕和黄桷树植冠损伤后的枝发生进行了研究.实验发现,切枝造成的植冠损伤对榕和黄桷树植株侧枝上的枝发生没有影响,不同损伤强度之间和不同损伤发生时间之间都不存在明显的差异.但是,植冠损伤后,榕和黄桷树植株主茎上的枝发生是位置依赖性的.植冠损伤不影响榕和黄桷树植株损伤处理后新生主茎段和具侧枝主茎段上的枝发生,却促进了损伤处理后裸露主茎段上的枝发生,并且裸露主茎段上枝发生的数量和密度随植冠损伤强度的升高而增大.此外,实验结果表明,秋季植冠损伤植株裸露主茎段上的枝发生要高于春季植冠损伤植株裸露主茎段上的枝发生;在榕和黄桷树的枝发生中,仅有裸露主茎段上增强的枝发生会促进植冠损伤植株对叶组织的生物量投资并有利于受损植株的恢复和生长.     

两种亚热带榕属乔木植冠损伤后的位置依赖性枝发生

三峡库区需要大量植株用于三峡工程所涉及的诸多建设(如铁路、公路、建筑)之后的植被恢复和绿化。由于具有优美树冠和耐瘠薄土壤的能力,榕(Ficus microcarpa L.)和黄桷树(F. virens Ait. var. sublanceolata (Miq.) Cornor)在三峡库区作为绿化和行道树木大量栽植。在三峡库区,这两种树种苗木的培育主要通过切枝扦插的方式进行。大量切枝损伤植株植冠并且使叶组织数量减少,对植株生长有很大影响。植株生长与植株的枝发生格局有很大关系,为明确植冠损伤对植株生长的可能影响,对榕和黄桷树植冠损伤后的枝发生进行了研究。实验发现,切枝造成的植冠损伤对榕和黄桷树植株侧枝上的枝发生没有影响,不同损伤强度之间和不同损伤发生时间之间都不存在明显的差异。但是,植冠损伤后,榕和黄桷树植株主茎上的枝发生是位置依赖性的。植冠损伤不影响榕和黄桷树植株损伤处理后新生主茎段和具侧枝主茎段上的枝发生,却促进了损伤处理后裸露主茎段上的枝发生,并且裸露主茎段上枝发生的数量和密度随植冠损伤强度的升高而增大。此外,实验结果表明, 秋季植冠损伤植株裸露主茎段上的枝发生要高于春季植冠损伤植株裸露主茎段上的枝发生; 在榕和黄桷树的枝发生中,仅有裸露主茎段上增强的枝发生会促进植冠损伤植株对叶组织的生物量投     

辽东栎芽库统计:芽的命运

植物体是一个构件集合体,植物的枝系伸展可由芽库出生率、死亡率的统计学过程来分析。在东灵山地区,应用随机枝取样法调查了辽东栎芽的命运,并对其与枝长、叶数、果数等的关系进行了统计分析。结果表明：（1）辽东栎的芽或保持休眠状态,或死亡后脱落,或分化为营养枝、生殖枝（包括雄花枝、雄花序、雌花枝和两花枝）等;（2）不同生境中芽的命运不同,生活在林窗中的幼树上的芽分化为具有生殖功能的枝条的比例显著高于郁闭林中的幼树,而与成熟个体接近;（3）芽的命运还受其它因子的影响,如上层枝条上、或叶数多的长枝上的芽分化为生殖枝的可能性大于其它的芽,另外还发现结实枝的枝长、枝上叶数都明显高于非结实枝。     

Growth parameters and resistance against Drechslera teres of spring barley (Hordeum vulgare L. cv. Scarlett) grown at elevated ozone and carbon dioxide concentrations

Spring barley ( Hordeum vulgare L. cv. Scarlett) was grown at two CO2 levels (400 vs. 700 ppm) combined with two ozone regimes (ambient vs. double ambient) in climate chambers for four weeks, beginning at seedling emergence. Elevated CO2 concentration significantly increased aboveground biomass, root biomass, and tiller number, whereas double ambient ozone significantly decreased these parameters. These ozone-induced reductions in growth parameters were strongly overridden by 700 ppm CO2. The elevated CO2 level increased C : N ratio of the leaf tissue and leaf starch content but decreased leaf protein levels. Exposure to double ambient ozone did not affect protein content and C : N ratio but dramatically increased leaf starch levels at 700 ppm CO2. Resistance against Drechslera teres (Sacc.) Shoemaker was increased in leaves grown at double ambient ozone but was less obvious at 700 ppm than at 400 ppm CO2. Constitutive activities of beta-1,3-glucanase and chitinase were significantly higher in leaves grown at double ambient ozone compared to ambient ozone levels. The sum of methanol-soluble and alkali-released cell wall-bound aromatic metabolites (i.e., C-glycosylflavones and several structurally unidentified metabolites) and lignin contents did not show any treatment-dependent differences.     

Structural characteristics and chemical composition of birch (Betula pendula) leaves are modified by increasing CO2 and ozone

Impacts of ozone and CO₂ enrichment, alone and in combination, on leaf anatomical and ultrastructural characteristics, nutrient status and cell wall chemistry in two European silver birch (Betula pendula Roth) clones were studied. The young soil‐growing trees were exposed in open‐top chambers over three growing seasons to 2 × ambient CO₂ and/or ozone concentrations in central Finland. The trees were measured for changes in altogether 35 variables of leaf structure, nutrients and cell wall chemistry of green leaves, and 20 of the measured variables were affected by CO₂ and/or O₃. Elevated CO₂ increased the size of chloroplasts and starch grains, number of mitochondria, P : N ratio, and contents of cell wall hemicellulose. Elevated CO₂ decreased the total leaf thickness, specific leaf area, concentrations of N, K, Cu, S and Fe, and contents of cell wall α‐cellulose, uronic acids, acid‐soluble lignin and acetone‐soluble extractives. Elevated ozone led to thinner leaves, higher palisade to spongy ratio, increased number of peroxisomes and mitochondria, reduced content of Mn, Zn, Cu, hemicellulose and uronic acids, and lower Mn : N and Zn : N ratios. In the combined exposure, interactions were antagonistic. Ultrastructural changes became more evident towards the end of the exposure. Young leaves were tolerant against ozone‐caused oxidative stress, whereas oxidative H₂O₂ accumulation was found in older leaves. CO₂ enrichment improved ozone tolerance not only through increased photosynthesis rates, but also through changes in cell wall chemistry (hemicellulose, in particular). However, nutrient imbalances due to ozone and/or CO₂ may predispose the trees to other biotic and abiotic stresses. Down‐regulation and up‐regulation of photosynthesis under elevated CO₂ through anatomical changes is discussed.     

Measuring and simulating crown respiration of Scots pine with increased temperature and carbon dioxide enrichment

Acclimation to elevated atmospheric carbon dioxide concentration and temperature of respiration by the foliage in the crown of Scots pine (Pinus sylvestris) trees is measured and modelled. Starting in 1996, individual 20-year-old trees were enclosed in chambers and exposed to either normal ambient conditions (CON), elevated CO2 concentration (EC), elevated temperature (ET) or a combination of EC and ET (ECT). Respiration of individual leaves within the crown was measured in 2000. To extrapolate the response of respiration of individual leaves to the whole crown, a multi-layer model was developed and used to predict daily and annual crown respiration, in which the crown structure and corresponding microclimate data were used as input. Respiration measurements showed that EC led to higher Q10 values (4.6%) relative to CON, but lower basal respiration rates at 20 degrees C [R1.d(20)] (-7.1%) during the main growth season (days 120-240), whereas ET and ECT both reduced Q10 (-12.0 and -9.8%, respectively) throughout the year but increased R1.d(20) (27.2 and 21.6%, respectively) during the period of no-growth, and slightly reduced R1.d(20) (-1.7 and -2.8%, respectively) during the main growth season. Model computations showed that annual crown respiration increased: (1) by 16% in EC, with 92% of this increase attributable to the increase in foliage area; (2) by 35% in ET, with 66% related to the increase in foliage area and 17% to the rise in ambient temperature; and (3) by 27% in the case of ECT, with 43% attributable to the increase in foliage area and 29% to the rise in ambient temperature. Changed respiration parameters for individual leaves, induced by treatments, made only a small contribution to the annual crown respiration compared with the increased foliage area. The effects of changes in crown architecture and nitrogen distribution, caused by treatments, on the daily and annual course of crown respiration are discussed.     

Variation among and within clones in formation of roots and shoots during micropropagation of Campanula isophylla

It is known that treatments enhancing shoot formation often suppress root formation and vice versa. It would be of interest to know if such negative correlations between formation of roots and shoots were also present among genetically different plants, given the same treatment, to ensure that selection for superior shoot formation would not lead to inadvertent decreases in the capacity for root formation. Height and dry weight of micropropagated shoot clusters and the numbers of shoots and roots were measured in 95 seedling clones. Within clones, shoot size was negatively correlated with number of shoots and positively correlated with number of roots. Among clones, however, the number of shoots was not correlated with the size of shoots, but positively correlated with the number of roots. While it is difficult to devise treatments that simultaneously optimize the initiation of roots and shoots, it is thus possible to select for fast-growing clones without compromising root formation.Abbreviations CM clonal means - DCM deviation from clonal means     

Seasonal variation in respiration of 1-year-old shoots of scots pine exposed to elevated carbon dioxide and temperature for 4 years

Sixteen 20-year-old Scots pine (Pinus sylvestris L.) trees growing in the field were enclosed for 4 years in environment-controlled chambers that maintained: (1) ambient conditions (CON); (2) elevated atmospheric CO2 concentration (ambient + 350 micro mol mol-1; EC); (3) elevated temperature (ambient +2-6 degrees C; ET); or (4) elevated CO2 and elevated temperature (ECT). The dark respiration rates of 1-year-old shoots, from which needles had been partly removed, were measured over the growing season in the fourth year. In all treatments, the temperature coefficient of respiration, Q10, changed with season, being smaller during the growing season than at other times. Respiration rate varied diurnally and seasonally with temperature, being highest around mid-summer and declining gradually thereafter. When measurements were made at the temperature of the chamber, respiration rates were reduced by the EC treatment relative to CON, but were increased by ET and ECT treatments. However, respiration rates at a reference temperature of 15 degrees C were reduced by ET and ECT treatments, reflecting a decreased capacity for respiration at warmer temperatures (negative acclimation). The interaction between season and treatment was not significant. Growth respiration did not differ between treatments, but maintenance respiration did, and the differences in mean daily respiration rate between the treatments were attributable to the maintenance component. We conclude that maintenance respiration should be considered when modelling respiratory responses to elevated CO2 and elevated temperature, and that increased atmospheric temperature is more important than increasing CO2 when assessing the carbon budget of pine forests under conditions of climate change.     

Shoot damage by Tomicus sp. (Coleoptera: Scolytidae) and effect on Pinus yunnanensis resistance to subsequent reproductive attacks in the stem

Abstract 1 In South‐western China, Yunnan pines Pinus yunnanensis, suffer considerable damage from an undescribed Tomicus sp. previously thought to be T. piniperda. 2 To assess the effect of shoot maturation feeding (during which an aggregation process appears to occur) on host resistance to attacks on the bole, the relationships between shoot damage, bole attack density and tree survival were studied. 3 Attack distribution in the crown and in the stem did not vary between killed and surviving trees, indicating that mortality is determined by the quantity of attacks. 4 The level of shoot damage and bole attack density were positively and linearly correlated. This can be explained by the fact that bole attacks are caused by beetles coming from the crown of the same tree. 5 A critical threshold of bole attack density (around 80 attacks/m²) above which trees die was observed. However, because attacks continue after this threshold is reached, the density of failed attacks on the killed trees was used as an estimator of the threshold density. It decreased when shoot damage increased. 6 The existence of a critical threshold of shoot damage (60% damaged shoots) was also demonstrated. Above this threshold, stem attack density was always sufficiently high to kill trees. 7 The results emphasize that concentration of shoot attacks is the main reason for the extensive tree damage observed in China. 8 A model of relationships between shoot and stem attacks is proposed, suggesting that management to reduce shoot attacks would protect trees from dying by both decreasing the number of bole attacks and raising the threshold for successful attack density on the bole to levels that could not be attained.     

Common allometric response of open-grown leader shoots to tree height in co-occurring deciduous broadleaved trees

Background and Aims

Morphology of crown shoots changes with tree height. The height of forest trees is usually correlated with the light environment and this makes it difficult to separate the effects of tree size and of light conditions on the morphological plasticity of crown shoots. This paper addresses the tree-height dependence of shoot traits under full-light conditions where a tree crown is not shaded by other crowns.

Methods

Focus is given to relationships between tree height and top-shoot traits, which include the shoot''s leaf-blades and non-leafy mass, its total leaf-blade area and the length and basal diameter of the shoot''s stem. We examine the allometric characteristics of open-grown current-year leader shoots at the tops of forest tree crowns up to 24 m high and quantify their responses to tree height in 13 co-occurring deciduous hardwood species in a cool-temperate forest in northern Japan.

Key Results

Dry mass allocated to leaf blades in a leader shoot increased with tree height in all 13 species. Specific leaf area decreased with tree height. Stem basal area was almost proportional to total leaf area in a leader shoot, where the proportionality constant did not depend on tree height, irrespective of species. Stem length for a given stem diameter decreased with tree height.

Conclusions

In the 13 species observed, height-dependent changes in allometry of leader shoots were convergent. This finding suggests that there is a common functional constraint in tree-height development. Under full-light conditions, leader shoots of tall trees naturally experience more severe water stress than those of short trees. We hypothesize that the height dependence of shoot allometry detected reflects an integrated response to height-associated water stress, which contributes to successful crown expansion and height gain.     

Shoot growth dynamics and size-dependent shoot fate of a clonal plant,Festuca rubra, in a mountain grassland

The relation of the within-season and between-season patterns of shoot growth were compared in a clonal grass with long-lived shoots,Festuca rubra, in a mown mountain grassland. The growth rate of shoot length from spring to summer in a year was almost constant for each shoot irrespective of spring shoot length each year. The annual shoot growth rate from spring to spring was negatively correlated with the shoot length in the first spring. Shoots of different length and age therefore tended to converge over time to a population of identical shoot size, suggesting an equalizing effect of growth pattern on size structure. Shoot size (shoot length and number of leaves) influenced the fates of shoots. Larger shoots showed an increased incidence of both flowering and formation of intravaginal daughter shoots and a decreased incidence of death in the subsequent time period. The fates of shoots were independent of their age. Although the negatively size-dependent springto-spring annual shoot growth rate acted to decrease shoot size variation, the remaining variation within the shoot population was still sufficient to generate different fates of shoots. These fates were not related to the previous life history of individual shoots. There was a significantly positive effect of the shoot size at initiation on its life expectancy. This was mainly attributable to the positively size-dependent survival rate of shoots in the early stage (<1 year old) of shoot life history. Later on (> 1 year old), shoot size had little effect on the survival rate of shoots. Once small young shoots have survived this early stage (< 1 year old) in life history, they can grow vigorously, little affected by competition regardless of shoot size, and converge to a stable size structure of shoots of similar size. Only shoot size in the early stage ( < 1 year old) of life history is important for the persistence of a shoot population.     

Shoot life span in relation to successional status in deciduous broad-leaved tree species in a temperate forest

In trees, leaf life span is closely related to successional status. Although leaves are attached to shoots, shoot life span has been insufficiently studied in the context of ecological systems. Interspecific variation in shoot survivorship was investigated over 27 months in 15 temperate hardwood tree species. Relationships between shoot architecture and shoot survival were also investigated. Shoot life span was shortest in early successional species, and longest in late successional species, in each of the families Betulaceae and Fagaceae. In Salicaceae, all of which were early successional species, shoot life span was longer in mountainous than in riparian species. Early successional or riparian species distributed longer shoots densely, even in proximal positions on mother shoots, resulting in mutual shading and consequent early and massive shoot shedding. By contrast, late successional or mountainous species concentrated shoots in distal positions, allowing shoots to receive equally favorable light, resulting in a longer life span. These results reveal close relationships between shoot life span and environmental resource availability or successional status and suggest a causal relationship between shoot shedding and shoot architecture.     

Modelling foliage characteristics in 3D tree crowns: influence on light interception and leaf irradiance

Because of the difficulty and time involved in making exhaustive measurements of the geometric parameters of large tree crowns, simplifying hypotheses are often used in 3D virtual plant modelling, but the effects on the radiation balance of each approximation are rarely assessed. Three hybrid walnut trees aged 7–9 years were digitized to analyse the effect of the crown geometric variables on light capture. The six studied variables were: (1) leaf area, (2) number of leaves per annual shoot, (3) position of leaves, (4) orientation of leaves, (5) leaflet inclination, and (6) lamina shape. For each variable, a sensitivity analysis compared a reference, based on observed values, with scenarios consisting of simplifying hypotheses. The total incident light intercepted during a bright day and the distributions of leaf irradiance were calculated using the Archimed radiative transfer model. Since some of the crown parameters were generated stochastically, the radiation simulations were repeated until results stabilised. Simplified models can be used to calculate with satisfactory results individual leaf area and number of leaves per shoot. Conversely, differentiating statistical distributions of individual leaf area between short and long shoots is more difficult and may generate errors up to 30%. Leaf clumping is a determining factor and requires correct grouping of leaves around the annual shoots bearing them. The effect of position of leaves along the shoot is less than 2%. Simple statistical distributions are adequate for representing leaf angle. Finally, the effect of specific leaf geometry is very important, but it can be approached using a limited number of representative leaf shapes.     

Testing a new hypothesis: plant vigor and phylloxera distribution on wild grape in Arizona

Summary Longer, meaning more vigorous, shoots of a wild grape clone (Vitis arizonica) were more susceptible to attack by second and third generations of leaf-galling grape phylloxera,Daktulopsphaira vitifoliae, as the growing season progressed. Although there was no significant difference in mean shoot length between attacked and unattacked shoots within a clone at the beginning of shoot elongation, attacked shoots were significantly longer than unattacked shoots when elongation had ceased (P<0.01). Also, long attacked shoots had a significantly greater population of phylloxera galls than short attacked shoots (P<0.01) as the season progressed. The phylloxera population on long shoots increased rapidly while the population on short shoots remained the same. Longer shoots also produced significantly more axillary shoots than shorter shoots as the season progressed (P<0.001), and the number of axillary shoots accounted for 66 percent of the variance in number of attacked leaves on a shoot. Experimental evidence showed that there was a significantly greater percentage of available leaves attacked on long shoots than on short shoots (P<0.05) and the leaves on long shoots generally had a greater number of galls per leaf. The relationship between shoot length and probability of attack was also tested by comparing shoots lengths of 10 attacked clones and 10 unattacked clones at a second location. Mean shoot lengths of attacked clones were significantly longer than mean shoot lengths of unattacked clones (P<0.05), and mean shoot lengths of attacked shoots within a clone were significantly longer than unattacked shoots (P<0.001). Longer shoot length accounted for 81 percent of the variance in probability of attack. The reason for this pattern of attack was that long shoots produced newly expanding leaves over a longer time during the growing season and multivoltine phylloxera require undifferentiated tissue to initiate gall formation. Patterns of attack within a shoot were characterized by an uneven distribution of galls among leaves. This was due to development time between generations and the current availability of undifferentiated tissue at times of colonization. This study supports the hypothesis that some herbivore species are favored more by vigorous plants than by stressed plants.     

火灾后兴安落叶松长短枝变化及其对生存的影响

研究了兴安落叶松的长短枝习性及火灾对其影响.结果表明,短枝在春季发叶迅速,而长枝的生长对兴安落叶松的枝条及树冠结构的形成起决定性作用,且在1个长枝上能形成10.5个短枝.中度火烧使兴安落叶松的树冠及枝条受到很大伤害,火烧可刺激1级侧枝上37%的短枝芽变成长枝,加速了火烧后兴安落叶松树冠结构的恢复,火后第3年兴安落叶松的短枝总量达到火前水平的98.46%.     

Shoot neoformation in clones of Fraxinus pennsylvanica in relation to genotype,site and pruning treatments

Summary Four clones of Fraxinus pennsylvanica var. subintegerrima (Vahl) Fern. were planted in replicated trials at two sites in Manitoba (Morden and Winnipeg) to investigate shoot growth and leaf neoformation in relation to genotype, environment and pruning treatment over a 3 year period. Significant differences were found among clones, years and sites for shoot length and numbers of neoformed leaves. Neoformation was highest shortly after transplanting and then declined. An increase in neoformation was evident following cold related winter injury or loss of terminal buds by late spring frosts. Pruning and terminal bud removal treatments both increased neoformed leaf production relative to control trees. The trees were able to quickly re-establish photosynthetic surface area after injury or treatment and neoformed leaf production was an important component in this recovery strategy. The capacity for neoformation also varied in relation to genotype but appeared to be very plastic, being affected by a wide variety of factors. In addition, the rate or pattern of change in amounts of neoformed leaves over time and locations was variable. Numbers of neoformed leaves increased with shoot length but variation in the relationship suggested that there were differences in internode length as well. Neoformed leaves were highest in the upper part of the crown indicating that there was differential allocation of resources within the crown.     

The effects of enhanced ozone and enhanced carbon dioxide concentrations on biomass, pigments and antioxidative enzymes in spruce needles (Picea abies L.)

During one growing period, 5-year-old spruce trees (Picea abies L., Karst.) were exposed in environmental chambers to elevated concentrations of carbon dioxide (750 cm³ m^?3) and ozone (008 cm³ m^?3) as single variables or in combination. Control concentrations of the gases were 350cm³ m^?3CO₂ and 0.02 cm³ m ^?3 ozone. To investigate whether an elevated CO₂ concentration can prevent adverse ozone effects by reducing oxidative stress, the activities of the protective enzymes superoxide dismutase, catalase and peroxidase were determined. Furthermore, shoot biomass, pigment and protein contents of two needle age classes were investigated. Ozone caused pigment reduction and visible injury in the previous year's needles and growth reduction in the current year's shoots. In the presence of elevated concentrations of ozone and CO₂, growth reduction in the current year's shoots was prevented, but emergence of visible damage in the previous year's needles was only delayed and pigment reduction was still found. Elevated concentrations of ozone or CO₂ as single variables caused a significant reduction in the activities of superoxide dismutase and catalase in the current year's needles. Minimum activities of superoxide dismutase and catalase and decreased peroxidase activities were found in both needle age classes from spruce trees grown at enhanced concentrations of both CO₂ and ozone. These results suggest a reduced tolerance to oxidative stress in spruce trees under conditions of elevated concentrations of both CO₂ and ozone.