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Carbon dioxide and water vapor exchange by young and old ponderosa pine ecosystems during a dry summer.

We investigated key factors controlling mass and energy exchange by a young (6-year-old) ponderosa pine (Pinus ponderosa Laws.) plantation on the west side of the Sierra Nevada Mountains and an old-growth ponderosa pine forest (mix of 45- and 250-year-old trees) on the east side of the Cascade Mountains, from June through September 1997. At both sites, we operated eddy covariance systems above the canopy to measure net ecosystem exchange of carbon dioxide and water vapor, and made concurrent meteorological and ecophysiological measurements. Our objective was to understand and compare the controls on ecosystem processes in these two forests. Precipitation is much higher in the young plantation than in the old-growth forest (1660 versus 550 mm year-1), although both forests experienced decreasing soil water availability and increasing vapor pressure deficits (D) as the summer of 1997 progressed. As a result, drought stress increased at both sites during this period, and changes in D strongly influenced ecosystem conductance and net carbon uptake. Ecosystem conductance for a given D was higher in the young pine plantation than in the old-growth forest, but decreased dramatically following several days of high D in late summer, possibly because of xylem cavitation. Net CO2 exchange generally decreased with conductance at both sites, although values were roughly twice as high at the young site. Simulations with the 3-PG model, which included the effect of tree age on fluxes, suggest that, during the fall through spring period, milder temperatures and ample water availability at the young site provide better conditions for photosynthesis than at the old pine site. Thus, over the long-term, the young site can carry more leaf area, and the climatic conditions between fall and spring offset the more severe limitations imposed by summer drought.

California↗

Ecosystem respiration in a young ponderosa pine plantation in the Sierra Nevada Mountains, California.

We estimated total ecosystem respiration from a ponderosa pine (Pinus ponderosa Dougl. ex Laws.) plantation in the Sierra Nevada Mountains near Georgetown, California, from June to October, 1998. We apportioned ecosystem respiration among heterotrophic, root, stem and foliage based on relationships for each component that considered microclimate and vegetation characteristics. We measured each respiration component at selected sampling points, and scaled the measurements up to the ecosystem based on modeled relationships. Over the study period, total mean ecosystem respiration was 5.7 +/- 1.3 mumol m-2 s-1 (based on daily mean), comprising about 67% from soil-surface CO2 efflux, 10% from stem and branch respiration and 23% from foliage respiration. Shrub leaves contributed about 24% to total foliage respiration, and current-year needles (1998 age class) accounted for 40% of total tree needle respiration. Root respiration accounted for 47% of soil-surface CO2 efflux. We conclude that ecosystem respiration can be estimated based on daily mean air and soil temperatures through exponential relationships with r2 values of 0.85 and 0.87, respectively. When based on both air and soil temperatures, about 91% of the variation in total ecosystem respiration could be explained by a linear regression.

Biomass↗

Blue wild-rye grass competition increases the effect of ozone on ponderosa pine seedlings.

Individual ponderosa pine (Pinus ponderosa Dougl. ex Laws.) seedlings were grown in mesocosms with three densities of blue wild-rye grass (Elymus glaucus Buckl.) (equivalent to 0, 32 or 88 plants m-2) to determine if the presence of a natural competitor alters the response of ponderosa pine seedlings to ozone. After 3 years of ozone exposure, grass presence reduced total ponderosa pine dry mass by nearly 50%, whereas ozone alone had no significant effect on ponderosa pine growth. The combination of ozone and grass further reduced needle, stem and branch dry mass significantly below that induced by grass competition alone. Root:shoot ratios increased in response to the combined grass and ozone treatments. Grass competition significantly reduced soluble sugar concentrations in all ponderosa pine tissue components examined. Starch concentrations were highly variable but did not differ significantly between treatments. Ozone significantly reduced soluble sugar concentrations in fine roots and stems. In the absence of grass, ozone-treated seedlings tended to have higher tissue N concentrations than controls. In the presence of grass, ozone-treated seedlings had lower N concentrations than controls, resulting in a significant interaction between these two stresses in 1- and 2-year-old needles. Needle C:N ratios decreased in response to grass competition, as a result of increased N concentration and no change in C concentration. The opposite response was observed in ozone-treated seedlings as a result of decreased N concentrations, indicating that ozone-treated seedlings were unable to take up or retain as much nitrogen when grown in the presence of grass. We conclude that ponderosa pine seedlings are more susceptible to ozone when grown in competition with blue wild-rye grass.

California↗

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

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

California↗

Effects of drought stress and high density stem inoculations with Leptographium wingfieldii on hydraulic properties of young Scots pine trees.

We examined drought-induced changes in susceptibility of potted Scots pine (Pinus sylvestris L.) trees to a bark-beetle associated fungus (Leptographium wingfieldii Morelet, from the bark beetle Tomicus piniperda L.). Five-year-old field-grown trees were transplanted to 50-l pots and grown for 1 year before the treatments were applied. Trees in the drought treatment were subjected to several successive, 3-week-long drought cycles, with predawn water potential dropping below -2 MPa at peak drought intensity. The experimental drought cycles were more severe than the natural drought episodes usually recorded in Scots pine stands. Trees were then mass-inoculated with L. wingfieldii at a density close to the critical threshold density of inoculations (400 m(-2)) above which tree resistance is overcome. Inoculation of well-watered trees resulted in induced reaction zones around the inoculation points and very limited damage (resinosis) in the sapwood. Drought alone had no long-lasting consequences on tree water relations, except for a decrease in hydraulic conductance in the youngest segments of the main stem. However, the combination of mass-inoculation and drought stress after inoculation resulted in a dramatic loss of stem hydraulic conductivity that was paralleled by conspicuous damage to the sapwood (resinosis, drying and blue staining). There was a close correlation between amount of visible sapwood damage and loss of hydraulic conductivity. The intensity of induced reactions in the phloem was unaffected by drought stress. We conclude that tree defence against L. wingfieldii is decreased during severe drought stress, resulting in changes in the spread and action of the fungus in the sapwood but not in the phloem.

Animals↗

Seasonal variation in nitrogen net uptake and root plasma membrane H+-ATPase activity of Scots pine seedlings as affected by nutrient availability.

We examined changes in nitrogen (N) net uptake and activity and amount of plasma membrane H+-ATPase (PM-ATPase) in roots of hydroponically cultured Scots pine (Pinus sylvestris L.) seedlings throughout a simulated second growing season. Seedlings were grown with low (0.25 mM N) or high (2.5 mM N) nutrient availability to determine whether root PM-ATPase is dependent on an external nutrient supply. Climatic conditions in the growth chamber simulated the mean growing season from May to mid-October in southern Finland. Root PM-ATPase activity varied considerably during the growing season and was higher in current-year roots than in previous-year roots. Total PM-ATPase activity of current-year roots was highest at the end of the growing season, whereas PM-ATPase activity per unit fresh mass of current-year roots and specific absorption rate of N were highest in mid-July and decreased at the end of the growing season. This indicates that the decrease in PM-ATPase activity per unit fresh mass of the roots at the end of the growing season was compensated by the increased size of the root system. Seasonal variation in PM-ATPase activity had no clear dependence on root zone temperature. The response of PM-ATPase to root zone temperature was dependent on the developmental stage of the seedling. High nutrient availability resulted in increased root PM-ATPase activity and an extended period of root growth in autumn.

Cell Membrane↗

Winter water relations of New England conifers and factors influencing their upper elevational limits. I. Measurements.

The upper elevational limits of tree species are thought to be controlled by abiotic factors such as temperature and the soil and atmospheric conditions affecting plant water status. We measured relative water contents (RWC), water potentials (Psi) and cuticular conductances (g(c)) of shoots of four conifer species-eastern hemlock (Tsuga canadensis (L.) Carr.), eastern white pine (Pinus strobus L.), red pine (P. resinosa Ait.) and red spruce (Picea rubens Sarg.)-during two winters on Mt. Ascutney, Vermont, USA. Some micrometeorological measurements are also reported. Eastern hemlock and white pine were studied near their upper elevational limits at a 640-m site, and red pine was studied near its upper elevational limit at 715 m. Red spruce was also studied at the 715-m site, which is in the middle of its elevational range on this mountain. There was no evidence of winter desiccation stress in any species. The observed distribution of seedlings suggested that the upper elevational limits on shade-intolerant eastern white pine and red pine are set by the absence of suitable seed beds after 100 years without fire. Eastern hemlock is able to reproduce in deep shade on organic substrates, but germination at high elevations may be restricted by low temperatures.

Altitude↗

Winter water relations of New England conifers and factors influencing their upper elevational limits. II. Modeling.

We used the water relations model, WINWAT, to model winter water relations of three conifer species-eastern hemlock (Tsuga canadensis (L.) Carr.), eastern white pine (Pinus strobus L.) and red pine (P. resinosa Ait.)-growing at their upper elevational limits on Mt. Ascutney, Vermont, USA, in the winters of 1997 and 1999. Modeled relative water contents remained above 60% in the two youngest foliar age classes of all three species during both winters, indicating that desiccation stress in winter is not responsible for setting the upper elevational limits of these species at this site under present climatic conditions. WINWAT indicated that winter water relations of these low-elevational species were sensitive to low relative humidity, which increased transpiration rates, and low temperatures, which inhibited recharge, but are much less sensitive to summer climate than in the case of subalpine conifers in Colorado. Our results indicate that summer and winter temperatures and relative humidities (or precipitation/potential evapotranspiration ratios) should be incorporated into climate change models designed to simulate future tree distributions.

Altitude↗

Effects of soil temperature on parameters of a coupled photosynthesis-stomatal conductance model.

To examine the effects of soil temperature on a coupled photosynthesis-stomatal conductance model, seedlings of trembling aspen (Populus tremuloides Michx.), jack pine (Pinus banksiana Lamb.), black spruce (Picea Mariana (Mill.) B.S.P.) and white spruce (Picea glauca (Moench) Voss) were exposed to soil temperatures ranging from 5 to 35 degrees C for 4 months. Light and CO(2) response curves of foliar gas exchange were measured for model parameterization. The effects of soil temperature on four key model parameters, V(cmax) (maximum rate of carboxylation), J(max) (maximum rate of electron transport), alpha (energy conversion efficiency or quantum efficiency of electron transport) and R(d) (daytime dark respiration), were modeled using two third-order polynomial equations and a modified Arrhenius equation. In all species, V(cmax) and J(max) increased with soil temperature up to an optimum, and then decreased with further increases in soil temperature. In the conifers, alpha showed a similar response to soil temperature as V(cmax) and J(max), but soil temperature had no significant effect on alpha in aspen. Soil temperature had no significant effect on R(d) in any species. The three equations described the relationships between soil temperature and the model parameters reasonably well, but performed best for V(cmax) and worst for alpha. No significant relationships were identified between soil temperature and the parameters of the stomatal conductance model.

Carbon Dioxide↗

Impacts of seasonal air and soil temperatures on photosynthesis in Scots pine trees.

Seasonal courses of light-saturated rate of net photosynthesis (A360) and stomatal conductance (gs) were examined in detached 1-year-old needles of Scots pine (Pinus sylvestris L.) from early April to mid-November. To evaluate the effects of soil frost and low soil temperatures on gas exchange, the extent and duration of soil frost, as well as the onset of soil warming, were manipulated in the field. During spring, early summer and autumn, the patterns of A360 and gs in needles from the control and warm-soil plots were generally strongly related to daily mean air temperatures and the frequency of severe frost. The warm-soil treatment had little effect on gas exchange, although mean soil temperature in the warm-soil plot was 3.8 degrees C higher than in the control plot during spring and summer, indicating that A360 and gs in needles from control trees were not limited by low soil temperature alone. In contrast, prolonged exposure to soil temperatures slightly above 0 degrees C severely restricted recovery of A360 and especially gs in needles from the cold-soil treatment during spring and early summer; however, full recovery of both A360 and gs occurred in late summer. We conclude that inhibition of A360 by low soil temperatures is related to both stomatal closure and effects on the biochemistry of photosynthesis, the relative importance of which appeared to vary during spring and early summer. During the autumn, soil temperatures as low as 8 degrees C did not affect either A360 or gs.

Carbon Dioxide↗

Canopy stomatal conductance and xylem sap abscisic acid (ABA) in mature Scots pine during a gradually imposed drought.

We investigated the effect of drought on canopy stomatal conductance (g(c)), and examined the hypothesis that g(c) is controlled by the chemical messenger abscisic acid (ABA) produced in roots. Beginning in November 1994, we subjected a mature stand of Scots pine (Pinus sylvestris L.) to an imposed 11-month drought. Control plots were maintained at average-season soil water content. Xylem sap was extracted from shoots at regular intervals from April to November 1995. Soil water, sap flow and leaf water potentials (predawn to dusk) were recorded at the same time. Canopy stomatal conductance was calculated from sap flow data and xylem sap ABA concentrations ([ABA(xyl)]) were measured by radioimmunoassay. Mean [ABA(xyl)] in control trees was 250 micromol m(-3). No diurnal variation in [ABA(xyl)] was detected. With soil drying, [ABA(xyl)] increased to a maximum in summer (600 micromol m(-3)), but decreased again toward autumn; however, no significant increase in ABA flux to the leaves occurred. A decline in g(c) was detected when volumetric soil water content declined below 0.12. The decline in g(c) could not have been mediated by increasing [ABA(xyl)] because stomatal closure appeared to precede any increase in [ABA(xyl)]. Peak sap flow velocity data were used to estimate delivery times for root-to-shoot signals in 15-m tall trees. Under normal field conditions, a signal would take 12 days to travel from the site of production (roots) to the presumed site of action (shoots). However, under drought conditions it may take a chemical signal in excess of 6 weeks. We conclude that a feedforward model of short-term stomatal response to soil drying, based solely on the action of a chemical messenger from the roots, is not applicable in mature conifer trees because signal transmission is too slow.

Abscisic Acid↗

Radial patterns of sap flow in woody stems of dominant and understory species: scaling errors associated with positioning of sensors.

We studied sap flow in dominant coniferous (Pinus sylvestris L.) and broadleaf (Populus canescens L.) species and in understory species (Prunus serotina Ehrh. and Rhododendron ponticum L.) by the heat field deformation (HFD) method. We attempted to identify possible errors arising during flow integration and scaling from single-point measurements to whole trees. Large systematic errors of -90 to 300% were found when it was assumed that sap flow was uniform over the sapwood depth. Therefore, we recommend that the radial sap flow pattern should be determined first using sensors with multiple measuring points along a stem radius followed by single-point measurements with sensors placed at a predetermined depth. Other significant errors occurred in the scaling procedure even when the sap flow radial pattern was known. These included errors associated with uncertainties in the positioning of sensors beneath the cambium (up to 15% per 1 mm error in estimated xylem depth), and differences in environmental conditions when the radial profile applied for integration was determined over the short term (up to 47% error). High temporal variation in the point-to-area correction factor along the xylem radius used for flow integration is also problematic. Compared with midday measurements, measurements of radial variation of sap flow in the morning and evening of sunny days minimized the influence of temporal variations on the point-to-area correction factor, which was especially pronounced in trees with a highly asymmetric sap flow radial pattern because of differences in functioning of the sapwood xylem layers. Positioning a single-point sensor at a depth with maximum sap flow is advantageous because of the high sensitivity of maximum sap flow to water stress conditions and changes in micro-climate, and because of the lower random errors associated with the positioning of a single-point sensor along the xylem radius.

Pinus↗

Radiation-use efficiency of a forest exposed to elevated concentrations of atmospheric carbon dioxide.

We compared radiation-use efficiency of growth (epsilon;), defined as rate of biomass accumulation per unit of absorbed photosynthetically active radiation, of forest plots exposed to ambient (approximately 360 micro l l-1) or elevated (approximately 560 micro l l-1) atmospheric CO2 concentration ([CO2]). Large plots (30-m diameter) in a loblolly pine (Pinus taeda L.) plantation, which contained several hardwood species in the understory, were fumigated with a free-air CO2 enrichment system. Biomass accumulation of the dominant loblolly pines was calculated from monthly measurements of tree growth and site-specific allometric equations. Depending on the species, leaf area index (L*) was estimated by three methods: optical, allometric and litterfall. Based on the relationship between tree height and diameter during the first 3 years of exposure, we conclude that elevated [CO2] did not alter the pattern of aboveground biomass allocation in loblolly pine. There was considerable variation in L* estimates by the different methods; total L* was 18-42% lower when estimated by the optical method compared with estimates from allometric calculations, and this discrepancy was reduced when optical measurements were corrected for the non-random distribution of loblolly pine foliage. The allometric + litterfall approach revealed a seasonal maximum total L* of 6.2-7.1 with about 1/3 of the total from hardwood foliage. Elevated [CO2] had only a slight effect on L* in the first 3 years of this study. Mean epsilon; (+/- SD), calculated for loblolly pine only, was 0.49 +/- 0.05 and 0.62 +/- 0.04 g MJ-1 for trees in the ambient and elevated [CO2] plots, respectively. The 27% increase in epsilon; in response to CO2 enrichment was caused primarily by the stimulation of biomass increment, as there was only a small effect of elevated [CO2] on L* during the initial years of fumigation. Long-term increases in atmospheric [CO2] can increase epsilon; in closed-canopy forests but the absolute magnitude and duration of this increase remain uncertain.

Biomass↗

Differences in delta13C and diameter growth among remnant Scots pine populations in Scotland.

Published data suggest that differences in wood cellulose carbon isotope composition (delta13C) and xylem ring width among natural populations of Scots pine in Scotland (Pinus sylvestris L.) are attributable to the persistence of palaeotypes of various post-glacial migratory origins. We assessed differences in wood cellulose delta13C and ring width among Scottish Scots pine populations grown in a clone bank and in natural stands at various locations in northern and central Scotland. Ring width and wood cellulose delta13C varied significantly among natural stands. Potential water deficit was positively correlated with wood cellulose delta13C and xylem ring width in the natural stands. Neither wood cellulose delta13C nor xylem ring width of clone bank trees correlated with any climate variables at the sites from which the trees originated, indicating little adaptation to climate for these traits. Xylem ring width showed a site x population interaction for the growth sites (i.e., natural stands versus clone bank), but wood cellulose delta13C did not. These results suggest that climate variation in Scotland has not resulted in significant genetic variation in wood cellulose delta13C or xylem ring width in post-glacial populations.

Carbon Isotopes↗

Leaf area dynamics of a boreal black spruce fire chronosequence.

Specific leaf area (SLA) and leaf area index (LAI) were estimated using site-specific allometric equations for a boreal black spruce (Picea mariana (Mill.) BSP) fire chronosequence in northern Manitoba, Canada. Stands ranged from 3 to 131 years in age and had soils that were categorized as well or poorly drained. The goals of the study were to: (i) measure SLA for the dominant tree and understory species of boreal black spruce-dominated stands, and examine the effect of various biophysical conditions on SLA; and (ii) examine leaf area dynamics of both understory and overstory for well- and poorly drained stands in the chronosequence. Overall, average SLA values for black spruce (n = 215), jack pine (Pinus banksiana Lamb., n = 72) and trembling aspen (Populus tremuloides Michx., n = 27) were 5.82 +/- 1.91, 5.76 +/- 1.91 and 17.42 +/- 2.21 m2 x kg-1, respectively. Foliage age, stand age, vertical position in the canopy and soil drainage had significant effects on SLA. Black spruce dominated overstory LAI in the older stands. Well-drained stands had significantly higher overstory LAI (P < 0.001), but lower understory LAI (P = 0.022), than poorly drained stands. Overstory LAI was negligible in the recent (3-12 years old) burn sites and highest in the 70-year-old burn site (6.8 and 3.0 in the well- and poorly drained stands, respectively), declining significantly (by 30-50%) from this peak in the oldest stands. Understory leaf area represented a significant portion (> 40%) of total leaf area in all stands except the oldest.

Ecosystem↗

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

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

Ecosystem↗

Ectomycorrhizal fungi and exogenous auxins influence root and mycorrhiza formation of Scots pine hypocotyl cuttings in vitro.

We studied the ability of the ectomycorrhizal (ECM) fungi, Pisolithus tinctorius (Pers.) Coker and Couch and Paxillus involutus (Batsch) Fr. (Strain H), to produce indole-3-acetic acid (IAA) and to affect the formation and growth of roots on Scots pine (Pinus sylvestris L.) hypocotyl cuttings in vitro. Effects of indole-3-butyric acid (IBA) and the auxin transport inhibitor, 2,3,5-triiodobenzoic acid (TIBA), on rooting and the cutting-fungus interaction were also studied. Both fungi produced IAA in the absence of exogenous tryptophan, but the mycelium and culture filtrate of Pisolithus tinctorius contained higher concentrations of free and conjugated IAA than the mycelium and culture filtrate of Paxillus involutus. Inoculation with either fungus or short-term application of culture filtrate of either fungus to the base of hypocotyl cuttings enhanced root formation. Inoculation with either fungus was even more effective in enhancing root formation than treatment of the hypocotyl bases with IBA. Fungal IAA production was not directly correlated with root formation, because rooting was enhanced more by Paxillus involutus than by Pisolithus tinctorius. This suggests that, in addition to IAA, other fungal components play an important role in root formation. Treatment with 5 microM TIBA increased the rooting percentage of non-inoculated cuttings, as well as of cuttings inoculated with Pisolithus tinctorius, perhaps as a result of accumulation of IAA at the cutting base. However, the marked reduction in growth of Pisolithus tinctorius in the presence of TIBA suggests that the effects of TIBA on rooting are complicated and not solely related to IAA metabolism. The high IAA-producer, Pisolithus tinctorius, formed mycorrhizas, and the IBA treatment increased mycorrhizal frequency in this species, whereas TIBA decreased it. Paxillus involutus did not form mycorrhizas, indicating that a low concentration of IAA together with other fungal components were sufficient to stimulate formation and growth of the roots, but not the formation of ECM symbiosis.

Basidiomycota↗

Impact of needle age on the response of respiration in Scots pine to long-term elevation of carbon dioxide concentration and temperature.

Sixteen 20-year-old Scots pine (Pinus sylvestris L.) trees growing in the field were enclosed in environment-controlled chambers that for 4 years maintained: (1) ambient conditions (CON); (2) elevated atmospheric carbon dioxide concentration [CO2] (ambient + 350 micromol mol-1; EC); (3) elevated temperature (ambient + 2-3 degrees C; ET); or (4) elevated [CO2] and temperature (EC+ET). Dark respiration rate, specific leaf area (SLA) and the concentrations of starch and soluble sugars in needles were measured in the fourth year. Respiration rates, on both an area and a mass basis, and SLA decreased in EC relative to CON, but increased in ET and EC+ET, regardless of needle age class. Starch and soluble sugar concentrations for a given needle age class increased in EC, but decreased slightly in ET and EC+ET. Respiration rates and SLA were highest in current-year needles in all treatments, whereas starch and soluble sugar concentrations were highest in 1-year-old needles. Relative to that of older needles, respiration of current-year needles was inhibited less by EC, but increased in response to ET and EC+ET. All treatments enhanced the difference in respiration between current-year and older needles relative to that in CON. Age had a greater effect on needle respiration than any of the treatments. There were no differences in carbohydrate concentration or SLA between needle age classes in response to any treatment. Relative to CON, the temperature coefficient (Q10) of respiration increased slightly in EC, regardless of age, but declined significantly in ET and EC+ET, indicating acclimation of respiration to temperature.

Carbohydrate Metabolism↗