Seasonal patterns of light-saturated photosynthesis and leaf conductance for mature and seedling Quercus rubra L. foliage: differential sensitivity to ozone exposure

Extrapolation of the effects of ozone on seedlings to large trees and forest stands is a common objective of current assessment activities, but few studies have examined whether seedlings are useful surrogates for understanding how mature trees respond to ozone. This two-year study utilized a replicated open-top chamber facility to test the effects of subambient, ambient and twice ambient ozone concentrations on light-saturated net photosynthesis (P(max)) and leaf conductance (g(l)) of leaves from mature trees and genetically related seedlings of northern red oak (Quercus rubra L.). Gas exchange measurements were collected four times during the 1992 and 1993 growing seasons. Both P(max) and g(l) of all foliage followed normal seasonal patterns of ontogeny, but mature tree foliage had greater P(max) and g(l) than seedling foliage at physiological maturity. At the end of the growing season, P(max) and g(l) of the mature tree foliage exposed to ambient ( approximately 80-100 ppm-h) and twice ambient ( approximately 150-190 ppm-h) exposures of ozone were reduced 25 and 50%, respectively, compared with the values for foliage in the subambient ozone treatment ( approximately 35 ppm-h). In seedling leaves, P(max) and g(l) were less affected by ozone exposure than in mature leaves. Extrapolations of the results of seedling exposure studies to foliar responses of mature forests without considering differences in foliar anatomy and stomatal response between juvenile and mature foliage may introduce large errors into projections of the response of mature trees to ozone.     

Effects of carbon dioxide, fertilization, and irrigation on photosynthetic capacity of loblolly pine trees

Branches of nine-year-old loblolly pine trees grown in a 2 x 2 factorial combination of fertilization and irrigation were exposed for 11 months to ambient, ambient + 175, or ambient + 350 micro mol mol(-1) CO(2). Rates of light-saturated net photosynthesis (A(max)), maximum stomatal conductance to water vapor (g(max)), and foliar nitrogen concentration (% dry mass) were assessed monthly from April 1993 until September 1993 on 1992 foliage (one-year-old) and from July 1993 to March 1994 on 1993 foliage (current-year). Rates of A(max) of foliage in the ambient + 175 CO(2) treatment and ambient + 350 were 32-47 and 83-91% greater, respectively, than that of foliage in the ambient CO(2) treatment. There was a statistically significant interaction between CO(2) treatment and fertilization or irrigation treatment on A(max) on only one measurement date for each age class of foliage. Light-saturated stomatal conductance to water vapor (g(max)) was significantly affected by CO(2) treatment on only four measurement dates. Light-saturated g(max) in winter was only 42% of summer g(max) even though soil water during winter was near field capacity and evaporative demand was low. Fertilization increased foliar N concentration by 30% over the study period when averaged across CO(2) treatments. During the study period, the ambient + 350 CO(2) treatment decreased average foliar N concentration of one-year-old foliage in the control, irrigated, fertilized and irrigated + fertilized plots by 5, 6.4, 9.6 and 11%, respectively, compared with one-year-old foliage in the corresponding ambient CO(2) treatments. The percent increase in A(max) due to CO(2) enrichment was similar in all irrigation and fertilization treatments and the effect persisted throughout the 11-month study period for both one-year-old and current-year foliage.     

Light environment alters ozone uptake per net photosynthetic rate in black cherry trees

Foliar ozone uptake rates of different-sized black cherry (Prunus serotina Ehrh.) trees were compared within a deciduous forest and adjacent openings in north-central Pennsylvania during one growing season. Study trees included open-grown seedlings and saplings, forest understory seedlings and saplings, and sunlit and shaded portions of mature canopy tree crowns. Instantaneous ozone uptake rates were highest in high-light environments primarily because of higher stomatal conductances. Low ozone uptake rates of seedlings and saplings in the forest understory could be attributed partially to lower average ambient ozone concentrations compared to the canopy and open environments. Among the tree size and light combinations tested, ozone uptake rates were highest in open-grown seedlings and lowest in forest-grown seedlings. Despite lower ozone uptake rates of foliage in shaded environments, ozone uptake per net photosynthesis of foliage in shaded environments was significantly higher than that of foliage in sunlit environments because of weaker coupling between net photosynthesis and stomatal conductance in shaded environments. The potential for greater ozone injury in shaded environments as a result of greater ozone uptake per net photosynthesis is consistent with previous reports of greater ozone injury in shaded foliage than in sunlit foliage.     

Canopy position and needle age affect photosynthetic response in field-grown Pinus radiata after five years of exposure to elevated carbon dioxide partial pressure

Photosynthesis of tree seedlings is generally enhanced during short-term exposure to elevated atmospheric CO2 partial pressure, but longer-term studies often indicate some degree of photosynthetic adjustment. We present physiological and biochemical evidence to explain observed long-term photosynthetic responses to elevated CO2 partial pressure as influenced by needle age and canopy position. We grew Pinus radiata D. Don. trees in open-top chambers for 5 years in sandy soil at ambient (36 Pa) and elevated (65 Pa) CO2 partial pressures. The trees were well watered and exposed to natural light and ambient temperature. In the fourth year of CO2 exposure (fall 1997), when foliage growth had ceased for the year, photosynthetic down-regulation was observed in 1-year-old needles, but not in current-year needles, suggesting a reduction in carbohydrate sink strength as a result of increasing needle age (Turnbull et al. 1998). In 5-year-old trees (spring 1997), when foliage expansion was occurring, photosynthetic down-regulation was not observed, reflecting significantly large sinks for carbohydrates throughout the tree. Net photosynthesis was stimulated by 79% in trees growing in elevated CO2 partial pressure, but there was no significant effect on photosynthetic capacity or Rubisco activity and concentration. Current-year needles were more responsive to elevated CO2 partial pressure than 1-year-old needles, exhibiting larger relative increases in net photosynthesis to elevated CO2 partial pressure (98 versus 64%). Lower canopy and upper canopy leaves exhibited similar relative responses to growth in elevated CO2 partial pressure. However, needles in the upper canopy exhibited higher net photosynthesis, photosynthetic capacity, and Rubisco activity and concentration than needles in the lower canopy. Given that the ratio of mature to juvenile foliage mass in the canopy will increase as trees mature, we suggest that trees may become less responsive to elevated CO2 partial pressure with increasing age. We conclude that tree response to elevated CO2 partial pressure is based primarily on sink strength and not on the duration of exposure.     

Rapid freezing induces winter injury symptomatology in red spruce foliage

Red spruce (Picea rubens Sarg.) suffers frequent and extensive injury to current-year foliage during the winter. Experimental freezing of red spruce foliage at cooling rates > 10 degrees C min(-1) induced visible symptomatology similar to natural winter injury at the branch, needle and cellular levels. Such damage was associated with a low-temperature exotherm near -10 to -12 degrees C, a loss in needle fluorescence, massive cellular disruption, foliar discoloration, and low needle survival. Susceptibility of individual trees to rapid freezing injury was associated with historical winter injury patterns and alterations in foliar nutrition. We conclude that anthropogenic deposition may alter the sensitivity of trees to winter injury caused by rapid temperature changes.     

Physiological responses of birch (Betula pendula) to ozone: a comparison between open-soil-grown trees exposed for six growing seasons and potted seedlings exposed for one season

Physiological responses of 4-year-old potted saplings of an O3-tolerant clone of Betula pendula Roth to short-term ozone (O3) exposure (one growing season) were compared with those of 6-year-old open-soil-grown trees of the same clone fumigated with O3 for six growing seasons. In the 2001 growing season, both groups of plants were exposed to ambient (control) and 1.6x ambient (elevated) O3 concentration under similar microclimatic conditions in a free air O3 exposure facility. Growth, net photosynthesis, stomatal conductance, stomatal density, visible foliar injury, starch and nutrient concentrations, bud formation and differences in O3 responses between lower, middle and upper sections of the canopy were determined. The potted saplings were unaffected by elevated O3 concentration, whereas the open-soil-grown trees showed a 3-38% reduction in shoot growth, a 22% reduction in number of overwintering buds, a 26-65% decrease in autumnal net photosynthesis, 30% and 20-23% reductions in starch and nitrogen concentrations of senescing leaves, respectively, and disturbances in stomatal conductance. The greater O3 sensitivity of open-soil-grown trees compared with potted saplings was a result of senescence-related physiological factors. First, a lower net photosynthesis to stomatal conductance ratio in open-soil-grown trees at the end of the season promoted O3 uptake and decreased photosynthetic gain, leading to the onset of visible foliar injuries. Second, decreased carbohydrate reserves may have resulted in deleterious carry-over effects arising from the reduced formation of over-wintering buds. Finally, the leaf-level O3 load was higher for open-soil-grown trees than for potted saplings because of slower leaf senescence in the trees. Thus, O3 sensitivity in European white birch increases with increasing exposure time and tree size.     

Carbon assimilation and nitrogen in needles of fertilized and unfertilized field-grown Scots pine at natural and elevated concentrations of CO2

Effects of elevated CO2 concentration ([CO2]) on carbon assimilation and needle biochemistry of fertilized and unfertilized 25-30-year-old Scots pine (Pinus sylvestris L.) trees were studied in a branch bag experiment set up in a naturally regenerated stand. In each tree, one branch was enclosed in a bag supplied with ambient [CO2] (360 micromol mol(-1)), a second branch was enclosed in a bag supplied with elevated [CO2] (680 micromol(-1)) and a control branch was left unbagged. The CO2 treatments were applied from April 15 to September 15, starting in 1993 for unfertilized trees and in 1994 for fertilized trees, which were treated with N in June 1994. Net photosynthesis, amount and activity of Rubisco, N, starch, C:N ratio and SLA of needles were measured during the growing season of 1995. Light-saturated net photosynthetic rates of 1-year-old and current-year shoots measured at ambient [CO2] were not affected by growth [CO2] or N fertilization. Elevated [CO2] reduced the amount and activity of Rubisco, and the relative proportion of Rubisco to soluble proteins and N in needles of unfertilized trees. Elevated [CO2] also reduced the chlorophyll concentration (fresh weight basis) of needles of unfertilized trees. Soluble protein concentration of needles was not affected by growth [CO2]. Elevated [CO2] decreased the Rubisco:chlorophyll ratio in unfertilized and fertilized trees. Starch concentration was significantly increased at elevated [CO2] only in 1-year-old needles of fertilized trees. Elevated [CO2] reduced needle N concentration on a dry weight or structural basis (dry weight minus starch) in unfertilized trees, resulting in an increase in needle C:N ratio. Fertilization had no effect on soluble protein, chlorophyll, Rubisco or N concentration of needles. The decrease in the relative proportions of Rubisco and N concentration in needles of unfertilized trees at elevated [CO2] indicates reallocation of N resources away from Rubisco to nonphotosynthetic processes in other plant parts. Acclimation occurred in a single branch exposed to high [CO2], despite the large sink of the tree. The responses of 1-year-old and current-year needles to elevation of growth [CO2] were similar.     

Carbon partitioning and allocation in northern red oak seedlings and mature trees in response to ozone

Northern red oak (Quercus rubra L.) seedlings and trees differ in their response to ozone. Previous work reported reductions in net photosynthesis, carboxylation efficiency and quantum yield of mature tree leaves, whereas seedling processes were unaffected by the same ozone exposure. To further characterize differences in ozone response between seedlings and mature trees, we examined carbon partitioning and allocation in 32-year-old trees and 4-year-old seedlings of northern red oak after exposure to subambient (seasonal SUM00 dose (sum of all hourly ozone exposures) = 31 ppm-h), ambient (SUM00 dose = 85 ppm-h) and twice ambient (SUM00 dose = 151 ppm-h) ozone concentrations for three growing seasons. For mature trees, ozone exposure decreased foliar starch partitioning, increased starch partitioning in branches and increased (14)C retention in leaves. In contrast, starch partitioning in leaves and branches, and foliar (14)C retention in seedlings were unaffected by ozone exposure, but soluble carbohydrate concentrations in coarse and fine roots of seedlings were reduced. Differences in carbohydrate demand between seedlings and mature trees may underlie the higher leaf ozone uptake rates and greater physiological response to ozone in mature northern red oak trees compared with seedlings.     

Effects of elevated CO(2) concentration and nutrition on net photosynthesis,stomatal conductance and needle respiration of field-grown Norway spruce trees

To study the effects of elevated CO(2) on gas exchange, nonstructural carbohydrate and nutrient concentrations in current-year foliage of 30-year-old Norway spruce (Picea abies (L.) Karst.) trees, branches were enclosed in ventilated, transparent plastic bags and flushed with ambient air (mean 370 &mgr;mol CO(2) mol(-1); control) or ambient air + 340 &mgr;mol CO(2) mol(-1) (elevated CO(2)) during two growing seasons. One branch bag was installed on each of 24 selected trees from control and fertilized plots. To reduce the effect of variation among trees, results from each treated branch were compared with those from a control branch on the same whorl of the same tree. Elevated CO(2) increased rates of light-saturated photosynthesis on average by 55% when measured at the treatment CO(2) concentration. The increase was larger in shoots with high needle nitrogen concentrations than in shoots with low needle nitrogen concentrations. However, shoots grown in elevated CO(2) showed a decrease in photosynthetic capacity compared with shoots grown in ambient CO(2). When measured at the internal CO(2) concentration of 200 &mgr;mol CO(2) mol(-1), photosynthetic rates of branches in the elevated CO(2) treatments were reduced by 8 to 32%. The elevated CO(2) treatment caused a 9 to 20% reduction in carboxylation efficiency and an 18% increase in respiration rates. In response to elevated CO(2), starch, fructose and glucose concentrations in the needles increased on average 33%, whereas concentrations of potassium, nitrogen, phosphorus, magnesium and boron decreased. Needle nitrogen concentrations explained 50-60% of the variation in photosynthesis and CO(2) acclimation was greater at low nitrogen concentrations than at high nitrogen concentrations. We conclude that the enhanced photosynthetic rates found in shoots exposed to elevated CO(2) increased carbohydrate concentrations, which may have a negative feedback on the photosynthetic apparatus and stimulate cyanide-resistant respiration. We also infer that the decrease in nutrient concentrations of needles exposed to elevated CO(2) was the result of retranslocation of nutrients to other parts of the branch or tree.     

Biochemical composition of loblolly pine reflects pollutant exposure

Under experimental conditions, the growth of loblolly pine (Pinus taeda L.) is often responsive to ozone at near-ambient concentrations. However, little is known of the biochemical changes associated with this or other pollutants. Loblolly pine seedlings in open-top chambers were exposed to combinations of ozone (sub-ambient, ambient, or twice-ambient), acidic precipitation (pH 3.8 or pH 5.2) and soil magnesium (0.15 or 0.32 microg g(-1) exchangeable Mg) for three growing seasons. The effects of these treatments were greater in foliage than in stems or roots. The largest treatment effect was a 50% decrease in the starch concentration of current-year foliage from the twice-ambient ozone treatment compared with current-year foliage from the sub-ambient ozone treatment. Responses to ozone were consistent with the hypothesis that ozone-induced growth reductions are associated with depletion of carbohydrate reserves resulting from injury compensation and repair processes or reduced carbon fixation or both. Addition of acidic precipitation, and to a small extent Mg, decreased sugar concentrations of tissues; however, this effect appeared to be mediated by nutrient addition rather than by acidity per se. Given the role of carbohydrates in plant resistance to environmental stress, the sensitivity of carbohydrates to experimental treatments demonstrates the potential for indirect effects of ozone, acidic precipitation, and soil properties on stress resistance. Noncarbohydrate constituents were largely unresponsive to the experimental treatments. These findings imply that tissue carbohydrate analysis may be useful for assessing the impacts of pollutants in forest ecosystems.     

Rapid freezing injury in red spruce: seasonal changes in sensitivity and effects of temperature range

On calm, cold days in winter, sun-exposed needles of red spruce (Picea rubens Sarg.) may warm 10 to 20 degrees C above ambient air temperature, and undergo rapid (>/= 1 degrees C min(-1)) fluctuations in temperature as light breezes or passing clouds alter the energy balance of the foliage. It has been proposed that the resulting rapid freeze-thaw cycles (freezing stress) cause a type of winter injury in montane red spruce that is characterized by necrosis of sun-exposed foliage. In autumn and winter, we monitored rapid freezing stress response of needle sections from 10 montane red spruce trees by subjecting needles to rapid freezing over the temperature span typically recorded in the field. In autumn, experimental rapid freezing stress produced severe injury only at temperatures considerably lower than expected for that time of year. In winter, rapid freezing caused occasional, moderate injury in fully hardened foliage of trees susceptible to both slow and rapid freezing. Seasonal changes in sensitivity to rapid and slow freezing were correlated, suggesting that environmental factors that are known to affect sensitivity to slow freezing may also affect sensitivity to rapid freezing. Experimental manipulation of the start and end temperatures of rapid freezing stress events showed that moderate to severe needle injury can occur in susceptible trees at temperature spans slightly more extreme than those typically recorded in the field. The extent of injury was similar at different starting temperatures if rapid freezing occurred over the same temperature span. Year-old foliage was consistently less sensitive to rapid freezing stress than current-year foliage, but some year-old foliage was damaged when the rapid freezing stress regime caused severe injury in current-year foliage. We conclude that rapid freeze-thaw cycles can explain light to moderate injury of current-year foliage, but they do not explain the more severe and widespread pattern of foliar damage that has occurred intermittently over at least the last 18 years.     

Mild ozone exposure alters (14)C dynamics in foliage of Pinus taeda L

To explore the physiological mechanisms underlying ozone-induced growth reductions in loblolly pine (Pinus taeda L.), seedlings were exposed to sub-ambient (charcoal-filtered), ambient or twice-ambient ozone in open-top chambers for three growing seasons. In the final year of exposure, current-year needle fascicles were labeled with (14)CO(2) and the incorporation of (14)C into biochemical fractions was followed for 48 hours. Irrespective of ozone treatment, losses of (14)C-assimilates from foliage to respiration and translocation were minimal during the first 3 hours, whereas more than 60% of the label was lost during the next 45 hours. Radiolabel in sugar decreased rapidly after a lag period, roughly paralleling the pattern of total (14)C loss. The amount of (14)C label in starch and lipids plus pigments remained constant throughout the 48-hour chase period, whereas the amount of (14)C label in other fractions showed a net decrease over the 48-hour chase period. Ozone treatments altered foliar carbon dynamics in two ways: (1) ozone exposure increased foliar (14)C retention up to 21% for the first 5 hours after labeling, but not thereafter, and (2) ozone exposure decreased partitioning of (14)C into starch and increased partitioning of (14)C into organic acids, residue, and lipids plus pigments, indicating an intensified partitioning of carbon to injury and repair processes. Both short-term carbon retention and diversion of carbon from storage compounds to repair processes are foliar mechanisms by which ozone exposure could decrease growth in loblolly pine seedlings.     

Influence of foliar N on foliar soluble sugars and starch of red spruce saplings exposed to ambient and elevated ozone

Red spruce (Picea rubens Sarg.) trees growing at high elevation in the northeastern United States have experienced decline in recent years but seedlings have proved to be relatively tolerant of a wide range of environmental stresses in controlled studies. One possible reason for the wide tolerance to stress in seedlings is their inherently large pool of carbohydrate reserves, which is available for maintenance during and regrowth after periods of stress. We tested for the effects of foliar N and exposure to ozone on foliar carbohydrate reserves of 20-year-old naturally regenerated saplings. The trees were maintained in native soil in 360-l containers for 5 years before the experiment. The year before the experiment, trees were fertilized with N,P,K to provide a population of trees from N deficient to N sufficient. As foliar N decreased below 0.9%, length of current-year shoots and specific needle area of current-year needles declined. Foliar N concentration was correlated with foliar sugar and starch concentrations, but relationships varied with time of year. Before bud break, foliar carbohydrates and N, in general, were positively correlated, and date of bud break was delayed in N-deficient trees. During active growth, foliar soluble sugars and N were positively correlated, but starch concentrations were negatively correlated with N. By late September, neither starch nor sugar concentration was correlated with N concentration. Ozone and foliar N concentrations did not interact to change foliar carbohydrate concentrations or shoot and needle growth in this relatively short-term study.     

Stomatal conductance alone does not explain the decline in foliar photosynthetic rates with increasing tree age and size in Picea abies and Pinus sylvestris

Foliar light-saturated net assimilation rates (A) generally decrease with increasing tree height (H) and tree age (Y), but it is unclear whether the decline in A is attributable to size- and age-related modifications in foliage morphology (needle dry mass per unit projected area; M(A)), nitrogen concentration, stomatal conductance to water vapor (G), or biochemical foliage potentials for photosynthesis (maximum carboxylase activity of Rubisco; V(cmax)). I studied the influences of H and Y on foliage structure and function in a data set consisting of 114 published studies reporting observations on more than 200 specimens of various height and age of Picea abies (L.) Karst. and Pinus sylvestris L. In this data set, foliar nitrogen concentrations were independent of H and Y, but net assimilation rates per unit needle dry mass (A(M)) decreased strongly with increasing H and Y. Although M(A) scaled positively with H and Y, net assimilation rates per unit area (A(A) = M(A) x A(M)) were strongly and negatively related to H, indicating that the structural adjustment of needles did not compensate for the decline in mass-based needle photosynthetic rates. A relevant determinant of tree height- and age-dependent modifications of A was the decrease in G. This led to lower needle intercellular CO2 concentrations and thereby to lower efficiency with which the biochemical photosynthetic apparatus functioned. However, V(cmax) per unit needle dry mass and area strongly decreased with increasing H, indicating that foliar photosynthetic potentials were lower in larger trees at a common intercellular CO2 concentration. Given the constancy of foliar nitrogen concentrations, but the large decline in apparent V(cmax) with tree size and age, I hypothesize that the decline in Vcmax results from increasing diffusive resistances between the needle intercellular air space and carboxylation sites in chloroplasts. Increased diffusive limitations may be the inevitable consequence of morphological adaptation (changes in M(A) and needle density) to greater water stress in needles of larger trees. Foliage structural and physiological variables were nonlinearly related to H and Y, possibly because of hyperbolic decreases in shoot hydraulic conductances with increasing tree height and age. Although H and Y were correlated, foliar characteristics were generally more strongly related to H than to Y, suggesting that increases in height rather than age are responsible for declines in foliar net assimilation capacities.     

Accretion, partitioning and sequestration of calcium and aluminum in red spruce foliage: implications for tree health

Calcium (Ca) is an essential macronutrient in plants and is an important component of many cellular structures and physiological processes as well as overall forest function. Aluminum (Al) in soil solution can inhibit Ca uptake by plants and disrupt many Ca-dependent metabolic and physiological processes of plants. The ratio of Ca to Al in soil solution can be an important indicator of forest health, especially on acid soils. We used sequential chemical extractions (water, acetic acid and hydrochloric acid) to assess the chemical availability of Ca and Al in foliage from mature red spruce (Picea rubens Sarg.) trees growing under ambient environmental conditions. In plants deficient in Ca and with intermediate total foliar Ca concentration ([Ca]), Ca preferentially accrued in labile and physiologically available forms (water- and acetic acid-extractable). In plants with total foliar [Ca] above a "sufficiency" threshold, Ca also accrued in a chemically sequestered form with low solubility (HCl-extractable), suggesting that Ca sequestration is an inducible process in response to excess foliar Ca. Because it has low solubility, it is likely that sequestered Ca is unavailable for Ca-dependent physiological processes. Immobilization of Al in foliage was related to Ca sequestration, suggesting that Ca sequestration may provide a passive mechanism for Al tolerance in the foliage of these trees. Aluminum immobilization was evident based on the ratio of HCl-extractable Al to the more labile (water- and acetic acid-extractable) forms of Al. Sufficient labile Ca combined with Al sequestration was associated with plant health, including enhanced foliar accretion of Mg and Mn, greater tree growth, enhanced foliar cold hardiness and reduced winter injury. These findings demonstrate that not all chemical forms of foliar Ca and Al are of equal physiological significance and underscore the importance of assessing the biologically significant element forms in biogeochemical research.     

Physiological responses of black spruce layers and planted seedlings to nutrient addition

We investigated effects of nutrient addition on several physiological characteristics of 60-cm-tall black spruce (Picea mariana Mill. B.S.P.) layers (i.e., rooted branches of overstory trees) and 20-cm-tall planted seedlings on a clear-cut, N-limited boreal site. After two growing seasons, current-year and one-year-old needles of fertilized trees (layers and seedlings combined) had higher net photosynthetic rates (A(n)) and maximum capacity of Rubisco for CO(2) fixation (V(max)) than unfertilized trees. One-year-old needles of fertilized trees had higher stomatal conductance (g(s)), higher water-use efficiency, and lower intercellular to ambient CO(2) ratio than unfertilized trees. Additionally, fertilized trees had higher predawn and midday shoot water potentials than unfertilized trees. Stomatal conductance of 1-year-old needles was 23% higher in seedlings than in layers, but there were no significant differences in g(s) of current-year needles between the regeneration types. For both needle age-classes, A(n) and V(max) of layers were 25 and 40% higher, respectively, than the corresponding values for seedlings. The higher values of A(n), V(max) and foliar N concentration of layers compared with seedlings after two growing seasons may be associated with the larger root systems of the layers compared with the transplanted seedlings.     

Sudden increase in atmospheric CO2 concentration reveals strong coupling between shoot carbon uptake and root nutrient uptake in young walnut trees

We studied the short-term (i.e., a few days) effect of a sudden increase in CO2 uptake by shoots on nutrient (NO3-, P ion, K+, Ca2+ and Mg2+) uptake by roots during vegetative growth of young walnut (Juglans nigra x J. major L.) trees. The increase in CO2 uptake was induced by a sudden increase in atmospheric CO2 concentration ([CO2]). Twelve 2-year-old trees were transplanted and grown in perlite-filled pots in a greenhouse. Rates of CO2 uptake and water loss by individual trees were determined by a branch bag method from 3 days before until 6 days after [CO2] was increased. Nutrient uptake rates were measured concurrently by a hydroponic recirculating nutrient solution system that provided non-limiting supplies of water and nutrients. Six control trees were kept in ambient [CO2] (360 ppm), and [CO2] was increased to 550 ppm for one set of three trees and to 800 ppm for another set of three trees. Before imposing the elevated [CO2] treatments, all trees exhibited similar daily water loss, CO2 uptake and nutrient uptake rates when expressed per unit leaf area to account for the tree size effect. Daily water loss rates were only slightly affected by elevated [CO2]. Carbon dioxide uptake rates greatly increased with increasing atmospheric [CO2], and nutrient uptake rates were proportional to CO2 uptake rates during the study period, except for P ion. Our results show that, despite the important carbon and nitrogen storage capacities previously observed in young walnut trees, nutrient uptake by roots is strongly coupled to carbon uptake by shoots over periods of a few days.     

Acid phosphatase activity and phosphorus nutrition in Scots pine needles

From March to October, acid phosphatase activity and phosphorus (P) concentration were measured in four needle age classes of the upper and lower crowns of fertilized and unfertilized Scots pine (Pinus sylvestris L.) trees. Negative correlations between acid phosphatase activity and P concentration were observed in current-year needles and in needles in the upper part of the crown, whereas there was a positive correlation between enzyme activity and P concentration in older needles and in needles in the lower part of the crown. In May and October, needles of all ages showed increased acid phosphatase activity. The most sensitive response of acid phosphatase activity to phosphate supply and phosphorus status of the whole tree was seen in current-year needles on the first whorl where a 300% increase in acid phosphatase activity was observed in response to a decrease in foliar P concentration of 1.7-1.8 mg P g(DW) (-1).     

Age effects on Norway spruce (Picea abies) susceptibility to ozone uptake: a novel approach relating stress avoidance to defense

Cumulative ozone (O3) uptake and O3 flux were related to physiological, morphological and biochemical characteristics of Norway spruce (Picea abies (L.) Karst.) trees of different ages. Under ambient CO2 conditions, photosynthetic capacity (Amax) declined in mature trees when cumulative O3 uptake into needles, which provides a measure of effective O3 dose, exceeded 21 mmol m-2 of total needle surface area. A comparable decline in Amax of seedlings occurred when cumulative O(3) uptake was only 4.5 mmol m-2. The threshold O3 flux causing a significant decline in Amax ranged between 2.14 and 2.45 nmol m-2 s-1 in mature trees and seedlings subjected to exposure periods of > or = 70 and > or = 23 days, respectively. The greater O3 sensitivity of young trees compared with mature trees was associated with needle morphology. Biomass of a 100-needle sample increased significantly with tree age, whereas a negative correlation was found for specific leaf area, these changes parallel those observed during differentiation from shade-type to sun-type needles with tree ontogeny. Age-dependent changes in leaf morphology were related to changes in detoxification capacity, with area-based concentrations of ascorbate increasing during tree ontogeny. These findings indicate that the extent of O3-induced injury is related to the ratio of potentially available antioxidants to O3 influx. Because this ratio, when calculated for ascorbate, increased with tree age, we conclude that the ratio may serve as an empirical basis for characterizing age-related differences in tree responses to O3.     

Effects of short-term ozone exposure and soil water availability on the carbon economy of juvenile Douglas-fir

Effects of ozone and soil water availability on partitioning and translocation of assimilates were studied in three-year-old Douglas-fir (Pseudotsuga menziesii [Mirb.] Franco) seedlings exposed, in separate experiments, to 0 and 106 or 0 and 514 micro g m(-3) ozone for 8 h day(-1) for 9 days. The dynamics of carbon from assimilated (14)CO(2) were followed. No interactions between ozone and soil water content were observed. Total net uptake of carbon was reduced by low soil water content, but was unaffected by ozone. Both ozone and low soil water content increased the amount of (14)C-photosynthates retained in the current-year needles. Total starch content in old and current-year needles was unaffected by ozone, but was reduced by low water availability. Translocation of carbon to the root-soil compartment was additively affected by ozone and low soil water content. The results suggest that dry periods in summer combined with high ozone concentrations cause the greatest reduction in the supply of carbon compounds to the root-soil compartment.