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Effects of temperature and nutrient availability on plasma membrane lipid composition in Scots pine roots during growth initiation.

We studied the effects of root zone temperature (RZT) and nutrient availability on free sterols and phospholipids in the plasma membrane (PM) and on PM-ATPase activity in roots of 1-year-old Scots pine (Pinus sylvestris L.) seedlings during growth initiation in the spring. Seedlings were grown for 6 weeks in hydroponic cultures with low (0.5 mM N; LN) or high (3 mM N; HN) nutrient availability. The root zone was subjected to slow warming (SW) and fast warming (FW) treatments while maintaining similar air temperatures in both treatments. Decreases in the amount of phospholipids and in the phospholipid/free sterol ratio, an increase in the degree of saturation of phospholipid fatty acids and changes in free sterol composition were observed during root growth initiation. Changes in lipid composition of the PM associated with the cold deacclimation process were detected at RZTs above 9 degrees C. Nutrient availability affected the lipid composition of the PM only when RZT was increased slowly. When RZT increased from 4 to 6 degrees C in the SW treatment, the degree of saturation of phospholipid fatty acids decreased, especially in HN seedlings. The sitosterol/stigmasterol ratio remained higher in HN seedlings than in LN seedlings. After an RZT of 9 degrees C had been reached in the SW treatment, HN caused increases in the saturation of phospholipid fatty acids and root PM-ATPase activity, and a decrease in the phospholipid/free sterol ratio. Possible effects of changes in PM lipid composition on root growth and PM-ATPase activity are discussed.

Adenosine Triphosphatases↗

A comparison of three approaches to modeling leaf gas exchange in annually drought-stressed ponderosa pine forests.

We tested, compared and modified three models of stomatal conductance at the leaf level in a forest ecosystem where drought stress is a major factor controlling forest productivity. The models were tested against 2 years (1999 and 2000) of leaf-level measurements on ponderosa pine (Pinus ponderosa Dougl. ex Laws.) growing in the Mediterranean climate of California, USA. The Ball, Woodrow and Berry (1987) (BWB) model was modified to account for soil water stress. Among the models, results of the modified BWB model were in the closest agreement with observations (r2 = 0.71). The Jarvis (1976) model showed systematic simulation errors related to vapor pressure deficit (r2 = 0.65). Results of the Williams, Rastetter, Fernandes et al. (1996) (SPA) model showed the poorest correlation with empirical data, but this model has only one calibration parameter (r2 = 0.60). Sensitivity analyses showed that, in all three models, predictions of stomatal conductance were most responsive to photosynthetically active radiation and soil water content. Stomatal conductance showed little sensitivity to vapor pressure deficit in the Jarvis model, whereas in both the BWB and SPA models, vapor pressure deficit (or relative humidity) was the third most important variable. Parameterization of the SPA model was in accordance with the parameterization of the modified BWB model, although the two models differ greatly. Measured and modeled results indicate that stomatal behavior is not water conservative during spring; however, during summer, when soil water content is low and vapor pressure deficit is high, stomatal conductance decreases and, according to the models, intrinsic water- use efficiency increases.

Climate↗

Age-related changes in ecosystem structure and function and effects on water and carbon exchange in ponderosa pine.

As forests age, their structure and productivity change, yet in some cases, annual rates of water loss remain unchanged. To identify mechanisms that might explain such observations, and to determine if widely different age classes of forests differ functionally, we examined young (Y, approximately 25 years), mature (M, approximately 90 years) and old (O, approximately 250 years) ponderosa pine (Pinus ponderosa Dougl. ex P. Laws.) stands growing in a drought-prone region of central Oregon. Although the stands differed in tree leaf area index (LAIT) (Y = 0.9, M = 2.8, O = 2.1), cumulative tree transpiration measured by sap flow did not differ substantially during the growing season (100-112 mm). Yet when water was readily available, transpiration per unit leaf area of the youngest trees was about three times that of M trees and five times that of O trees. These patterns resulted from a nearly sixfold difference in leaf specific conductance (KL) between the youngest and oldest trees. At the time of maximum transpiration in the Y stand in May-June, gross carbon uptake (gross ecosystem production, GEP) was similar for Y and O stands despite an almost twofold difference in stand leaf area index (LAIS). However, the higher rate of water use by Y trees was not sustainable in the drought-prone environment, and between spring and late summer, KL of Y trees declined fivefold compared with a nearly twofold decline for M trees and a < 30% reduction in O trees. Because the Y stand contained a significant shrub understory and more exposed soil, there was no appreciable difference in mean daily latent energy fluxes between the Y stand and the older stands as measured by the eddy-covariance technique. These patterns resulted in 60 to 85% higher seasonal GEP and 55 to 65% higher water-use efficiency at the M and O stands compared with the Y stand.

Carbon Dioxide↗

Mist, substrate water potential and cutting water potential influence rooting of stem cuttings of loblolly pine.

We investigated the influence of cutting water potential (Psicut) on rooting of juvenile hardwood (dormant) and softwood (succulent) stem cuttings of loblolly pine (Pinus taeda L.) propagated under varying substrate water potentials (Psisub) and volumes of mist application. Mist treatment and Psisub contributed to the Psicut of unrooted stem cuttings. When Psisub was held constant across mist treatments, mist treatment contributed strongly to Psicut. Substrate water potential affected rooting percentage when mist treatment was sub-optimal or excessive, otherwise mist treatment had a stronger effect than Psisub on rooting percentage. Cuttings rooted best when subjected to moderate cutting water potentials (-0.5 to -1.2 MPa) during the initial 4 or 5 weeks of the rooting period. Cuttings experiencing either severe water deficit or no water deficit rooted poorly. We conclude that the rooting environment should impose a moderate water stress on loblolly pine stem cuttings to achieve optimum rooting.

Atmosphere↗

Converging patterns of uptake and hydraulic redistribution of soil water in contrasting woody vegetation types.

We used concurrent measurements of soil water content and soil water potential (Psi(soil)) to assess the effects of Psi(soil) on uptake and hydraulic redistribution (HR) of soil water by roots during seasonal drought cycles at six sites characterized by differences in the types and amounts of woody vegetation and in climate. The six sites included a semi-arid old-growth ponderosa pine (Pinus ponderosa Dougl. ex P. Laws & C. Laws) forest, a moist old-growth Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) forest, a 24-year-old Douglas-fir forest and three Brazilian savanna sites differing in tree density. At all of the sites, HR was confined largely to the upper 60 cm of soil. There was a common threshold relationship between the relative magnitude of HR and Psi(soil) among the six study sites. Below a threshold Psi(soil) of about -0.4 MPa, overnight recharge of soil water storage increased sharply, and reached a maximum value of 80-90% over a range of Psi(soil) from ~ -1.2 to -1.5 MPa. Although amounts of water hydraulically redistributed to the upper 60 cm of soil were relatively small (0 to 0.4 mm day(-1)), they greatly reduced the rates of seasonal decline in Psi(soil). The effectiveness of HR in delaying soil drying diminished with increasing sapwood area per ground area. The relationship between soil water utilization and Psi(soil) in the 20-60-cm layer was nearly identical for all six sites. Soil water utilization varied with a surrogate measure of rhizosphere conductance in a similar manner at all six sites. The similarities in relationships between Psi(soil) and HR, soil water utilization and relative rhizosphere conductance among the six sites, suggests that, despite probable differences in maximum rooting depth and density, there was a convergence in biophysical controls on soil water utilization and redistribution in the upper soil layers where the density of finer roots is greatest.

Climate↗

Stomata open at night in pole-sized and mature ponderosa pine: implications for O3 exposure metrics.

Ponderosa pine (Pinus ponderosa Dougl. ex Laws.) is widely distributed in the western USA. We report the lack of stomatal closure at night in early summer for ponderosa pine at two of three sites investigated. Trees at a third site with lower nitrogen dioxide and nitric acid exposure, but greater drought stress, had slightly open stomata at night in early summer but closed stomata at night for the rest of the summer. The three sites had similar background ozone exposure during the summer of measurement (2001). Nighttime stomatal conductance (gs) ranged from one tenth to one fifth that of maximum daytime values. In general, pole-sized trees (< 40 years old) had greater nighttime gs than mature trees (> 250 years old). In late summer, nighttime gs was low (< 3.0 mmol H2O m(-2) s(-1)) for both tree size classes at all sites. Measurable nighttime gs has also been reported in other conifers, but the values we observed were higher. In June, nighttime ozone (O3) uptake accounted for 9, 5 and 3% of the total daily O3 uptake of pole-sized trees from west to east across the San Bernardino Mountains. In late summer, O3 uptake at night was < 2% of diel uptake at all sites. Nocturnal O3 uptake may contribute to greater oxidant injury development, especially in pole-sized trees in early summer.

Ozone↗

Boron retranslocation in Scots pine and Norway spruce.

We previously traced 10B-enriched boric acid from shoots to roots to demonstrate the translocation of boron (B) in Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) seedlings. To gain a more detailed understanding of B translocation, we sought: (1) to demonstrate B retranslocation directly, by showing that foliar-applied 10B is located in the new growth after dormancy; and (2) to assess whether shoot-applied B affects growth in the long term. We applied 10B-enriched boric acid to needles of Scots pine and Norway spruce seedlings. After a dormancy period and 9 weeks of growth, small but significant increases in the 10B isotope were found in the new stem and needles of both species. In Scots pine, the total B concentration of the new stem was also increased. Both species contained polyols, particularly pinitol and inositol. Boron-polyol complexes may provide a mechanism for mobilizing B in these species. To determine the long-term effects of applied B, seedlings were grown for two growing seasons after the application of 10B to shoots. In Norway spruce, the proportion of 10B in the root systems and current needles of the harvest year was slightly higher than in the controls, and in Scots pine root systems, marginally so. The B treatment had no effect on growth of Norway spruce seedlings. In Scots pine seedlings, the B treatment caused a 33% increase in total dry mass and significantly increased the number of side branches.

Boron↗

Winter photoinhibition in the field involves different processes in four co-occurring Mediterranean tree species.

Photoinhibition was examined in four co-occurring Mediterranean evergreen tree species during two consecutive winters. In response to low temperatures and saturating light, Juniperus phoenicea L., Pinus halepensis Mill., Quercus coccifera L. and Q. ilex ssp. ballota (Desf.) Samp. exhibited marked chronic photoinhibition, indicated by low predawn maximal photochemical efficiency of photosystem II (PSII) (Fv/Fm). Low Fv/Fm values were correlated with high concentrations of xanthophyll cycle components (VAZ) and with the maintenance of high concentrations of zeaxanthin overnight (DPSpd). In all species, however, chronic photoinhibition was enhanced as the winter progressed in the absence of changes in DPSpd, suggesting cumulative damage toward the end of winter. Photoinhibition differed among species: P. halepensis always displayed significantly higher Fv/Fm values; and Q. coccifera had the lowest Fv/Fm values, showing a high sensitivity to the combination of high light and low temperatures. Differences among species were not fully explained by differences in the xanthophyll pool or its de-epoxidation state. Chronic photoinhibition overlapped with a dynamic photoinhibition as shown by the low values of photochemical efficiency of the open reaction centers of PSII at midday. Winter photoprotective strategies differed among species and may involve photoprotective mechanisms in addition to those associated with xanthophylls. The observed species-specific differences matched results obtained for the same species in summer; however, comparison of the two seasons suggests that the higher VAZ concentration observed in winter has an additional structural photoprotective role.

Chlorophyll↗

Light sources with different spectra affect root and mycorrhiza formation in Scots pine in vitro.

We studied the effects of broad-spectrum light quality on the interaction between the ectomycorrhizal fungus Pisolithus tinctorius (Pers.) Coker and Couch and Scots pine (Pinus sylvestris L.) seedlings and hypocotyl cuttings cultured in vitro. The light sources were cool white (CW), warm white (WW) and red-rich daylight (RD) fluorescent lamps. Inoculation with P. tinctorius enhanced adventitious root formation of the cuttings in all light treatments. Rooting of the inoculated cuttings was highest in WW light (89%), followed by CW (73%) and RD light (66%). During 6 weeks of in vitro culture, rooted cuttings formed only a few lateral roots. The fungus grew over lateral roots, but the Hartig net was absent in all light treatments. In non-inoculated cuttings, neither root formation nor subsequent root growth was affected by light quality. In the seedling experiment, inoculation in the WW treatment resulted in a significantly (P < 0.05) greater number of lateral roots than inoculation in the RD treatment. The percentage of lateral roots covered with fungal hyphae was also highest in WW light (62%), followed by CW (50%) and RD (27%) light. A similar pattern was observed in the intensity of Hartig net formation. We conclude that effects of broad-spectrum light quality on the ectomycorrhizal fungus-root interaction are dependent on the developmental stage of the root.

Basidiomycota↗

Carbon budget for Scots pine trees: effects of size, competition and site fertility on growth allocation and production.

Time series of carbon fluxes in individual Scots pine (Pinus sylvestris L.) trees were constructed based on biomass measurements and information about component-specific turnover and respiration rates. Foliage, branch, stem sapwood, heartwood and bark components of aboveground biomass were measured in 117 trees sampled from 17 stands varying in age, density and site fertility. A subsample of 32 trees was measured for belowground biomass excluding fine roots. Biomass of fine roots was estimated from the results of an earlier study. Statistical models were constructed to predict dry mass (DW) of components from tree height and basal area, and time derivatives of these models were used to estimate biomass increments from height growth and basal area growth. Biomass growth (G) was estimated by adding estimated biomass turnover rates to increments, and gross photosynthetic production (P) was estimated by adding estimated component respiration rates to growth. The method, which predicts the time course of G, P and biomass increment in individual trees as functions of height growth and basal area growth, was applied to eight example trees representing different dominance positions and site fertilities. Estimated G and P of the example trees varied with competition, site fertility and tree height, reaching maximum values of 22 and 43 kg(DW) year(-1), respectively. The site types did not show marked differences in productivity of trees of the same height, although height growth was greater on the fertile site. The G:P ratio decreased with tree height from 65 to 45%. Growth allocation to needles and branches increased with increasing dominance, whereas growth allocation to the stem decreased. Growth allocation to branches decreased and growth allocation to coarse roots increased with increasing tree size. Trees at the poor site allocated 49% more to fine roots than trees at the fertile site. The belowground parts accounted for 25 to 55% of annual G, increasing with tree size and decreasing with site fertility. Annual G and P per unit needle mass varied over the ranges 1.9-2.4 and 3.5-4.0 kg(DW) kg(-1), respectively. The relationship between P and needle mass in the example trees was linear and relatively independent of competition, site fertility and age.

Biomass↗

Respiratory responses of Scots pine stems to 5 years of exposure to elevated CO2 concentration and temperature.

Stem respiration in 20-year-old Scots pine (Pinus sylvestris L.) trees was examined following 5 years of exposure to ambient conditions (CON), elevated atmospheric carbon dioxide concentration ([CO2]) (ambient + 350 micromol mol(-1), (EC)), elevated temperature (ambient + 2-6 degrees C, (ET)) or a combination of elevated [CO2] and elevated temperature (ECT). Stem respiration varied seasonally regardless of the treatment and displayed a similar trend to temperature, with maximum rates occurring around Day 190 in summer and minimum rates in winter. Respiration normalized to 15 degrees C (R15) was higher in the growing season than in the non-growing season, whereas the temperature coefficient (Q10) was lower in the growing season. Annually averaged R15 was 0.36, 0.43, 0.40 and 0.44 micromol m(-2) s(-1) under CON, EC, ET and ECT conditions, respectively, whereas the corresponding values for total stem respiration were 6.55, 7.69, 7.50 and 7.90 mol m(-2) year(-1). The EC, ET and ECT treatments increased R15 by 18, 11 and 22%, respectively, relative to CON, and increased the modeled annual total stem respiration by 18, 15 and 21%. The increase in modeled annual stem respiration under EC and ECT conditions was caused mainly by higher maintenance respiration (22 and 25%, respectively, whereas the increase in growth respiration was 9 and 12%). Growth respiration was unaltered by ET. The treatments did not significantly affect the respiratory response to stem temperature; the mean Q10 value was 2.04, 2.10, 1.99 and 2.12 in the CON, EC, ET and ECT treatments, respectively. It is suggested that the increase in stem respiration was partly a result of the increased growth rate. We conclude that elevated [CO2] increased the maintenance component of respiration more than the growth component.

Carbon Dioxide↗

Forest thinning and soil respiration in a ponderosa pine plantation in the Sierra Nevada.

Soil respiration is controlled by soil temperature, soil water, fine roots, microbial activity, and soil physical and chemical properties. Forest thinning changes soil temperature, soil water content, and root density and activity, and thus changes soil respiration. We measured soil respiration monthly and soil temperature and volumetric soil water continuously in a young ponderosa pine (Pinus ponderosa Dougl. ex P. Laws. & C. Laws.) plantation in the Sierra Nevada Mountains in California from June 1998 to May 2000 (before a thinning that removed 30% of the biomass), and from May to December 2001 (after thinning). Thinning increased the spatial homogeneity of soil temperature and respiration. We conducted a multivariate analysis with two independent variables of soil temperature and water and a categorical variable representing the thinning event to simulate soil respiration and assess the effect of thinning. Thinning did not change the sensitivity of soil respiration to temperature or to water, but decreased total soil respiration by 13% at a given temperature and water content. This decrease in soil respiration was likely associated with the decrease in root density after thinning. With a model driven by continuous soil temperature and water time series, we estimated that total soil respiration was 948, 949 and 831 g C m(-2) year(-1) in the years 1999, 2000 and 2001, respectively. Although thinning reduced soil respiration at a given temperature and water content, because of natural climate variability and the thinning effect on soil temperature and water, actual cumulative soil respiration showed no clear trend following thinning. We conclude that the effect of forest thinning on soil respiration is the combined result of a decrease in root respiration, an increase in soil organic matter, and changes in soil temperature and water due to both thinning and interannual climate variability.

California↗

Scots pine responses to elevated temperature and carbon dioxide concentration: growth and wood properties.

Growth and wood properties of 20-year-old Scots pine (Pinus sylvestris L.) trees were studied for 6 years in 16 closed chambers providing a factorial combination of two temperature regimes (ambient and elevated) and two carbon dioxide concentrations ([CO2]) (ambient and twice ambient). The elevation of temperature corresponded to the predicted effect at the site of a doubling in atmospheric [CO2]. Annual height and radial growth and wood properties were analyzed during 1997-2002. Physical wood properties analyzed included early- and latewood widths and their proportions, intra-ring wood densities, early- and latewood density and mean fiber length. Chemical wood properties analyzed included concentrations of acetone-soluble extractives, lignin, cellulose and hemicellulose. There were no significant treatment effects on height growth during the 6-year study. Elevated [CO2] increased ring width by 66 and 47% at ambient and elevated temperatures, respectively. At ambient [CO2], elevated temperature increased ring width by 19%. Increased ring width in response to elevated [CO2] resulted from increases in both early- and latewood width; however, there was no effect of the treatments on early- and latewood proportions. Mean wood density, earlywood density and fiber length increased in response to elevated temperature. The chemical composition of wood was affected by elevated [CO2], which reduced the cellulose concentration, and by elevated temperature, which reduced the concentration of acetone-soluble extractives. Thus, over the 6-year period, radial growth was significantly increased by elevated [CO2], and some wood properties were significantly affected by elevated temperature or elevated [CO2], or both, indicating that climate change may affect the material properties of wood.

Carbon Dioxide↗

Element accumulation patterns of deciduous and evergreen tree seedlings on acid soils: implications for sensitivity to manganese toxicity.

Foliar nutrient imbalances, including the hyperaccumulation of manganese (Mn), are correlated with symptoms of declining health in sensitive tree species growing on acidic forest soils. The objectives of this study were to: (1) compare foliar nutrient accumulation patterns of six deciduous (sugar maple (Acer saccharum Marsh.), red maple (Acer rubrum L.), red oak (Quercus rubra L.), white oak (Quercus alba L.), black cherry (Prunus serotina Ehrh.) and white ash (Fraxinus americana L.)) and three evergreen (eastern hemlock (Tsuga canadensis L.), white pine (Pinus strobus L.) and white spruce (Picea glauca (Moench) Voss.)) tree species growing on acidic forest soils; and (2) examine how leaf phenology and other traits that distinguish evergreen and deciduous tree species influence foliar Mn accumulation rates and sensitivity to excess Mn. For the first objective, leaf samples of seedlings from five acidic, non-glaciated field sites on Pennsylvania's Allegheny Plateau were collected and analyzed for leaf element concentrations. In a second study, we examined growth and photosynthetic responses of seedlings exposed to excess Mn in sand culture. In field samples, Mn in deciduous foliage hyperaccumulated to concentrations more than twice as high as those found in evergreen needles. Among species, sugar maple was the most sensitive to excess Mn based on growth and photosynthetic measurements. Photosynthesis in red maple and red oak was also sensitive to excess Mn, whereas white oak, black cherry, white ash and the three evergreen species were tolerant of excess Mn. Among the nine species, relative rates of photosynthesis were negatively correlated with foliar Mn concentrations, suggesting that photosynthetic sensitivity to Mn is a function of its rate of accumulation in seedling foliage.

Acer↗

Modeling tree water deficit from microclimate: an approach to quantifying drought stress.

Tree water deficit estimated by measuring water-related changes in stem radius (DeltaW) was compared with tree water deficit estimated from the output of a simple, physiologically reasonable model (DeltaWE), with soil water potential (Psisoil) and atmospheric vapor pressure deficit (VPD) as inputs. Values of DeltaW were determined by monitoring stem radius changes with dendrometers and detrending the results for growth. We followed changes in DeltaW and DeltaWE in Pinus sylvestris L. and Quercus pubescens Willd. over 2 years at a dry site (2001-2002; Salgesch, Wallis) and in Picea abies (L.) Karst. for 1 year at a wet site (1998; Davos, Graubuenden) in the Swiss Alps. The seasonal courses of DeltaW in deciduous species and in conifers at the same site were similar and could be largely explained by variation in DeltaWE. This finding strongly suggests that DeltaW, despite the known species-specific differences in stomatal response to microclimate, is mainly explained by a combination of atmospheric and soil conditions. Consequently, we concluded that trees are unable to maintain any particular DeltaW. Either Psisoil or VPD alone provided poorer estimates of DeltaW than a model incorporating both factors. As a first approximation of DeltaWE, Psisoil can be weighted so that the negative mean Psisoil reaches 65 to 75% of the positive mean daytime VPD over a season (Q. pubescens: approximately 65%, P. abies: approximately 70%, P. sylvestris: approximately 75%). The differences in DeltaW among species can be partially explained by a different weighting of Psisoil against VPD. The DeltaW of P. sylvestris was more dependent on Psisoil than that of Q. pubescens, but less than that of P. abies, and was less dependent on VPD than that of P. abies and Q. pubescens. The model worked well for P. abies at the wet site and for Q. pubescens and P. sylvestris at the dry site, and may be useful for estimating water deficit in other tree species.

Climate↗

Seasonal variations in location and population structure of endophytes in buds of Scots pine.

We studied the location and distribution of a bacterial isolate, a Mycobacterium sp., in buds of Scots pine (Pinus sylvestris L.). Using a probe specific for the 16S rRNA of the Mycobacterium sp., the bacterium was found by in situ hybridization in the meristematic tissues of 40% of all bud samples examined. Because we had previously found other bacterial and fungal endophytes in the meristematic tissues of Scots pine buds, we studied their occurrence in buds during shoot development and dormancy. Using probes targeted to the 16S or 18S rRNA of the endophytes Mycobacterium sp., Methylobacterium spp., Pseudomonas spp. and Rhodotorula minuta, endophytes were found in association with growing tissues, with Methylobacterium spp. being the dominant species. Endophytes were detected in abundance before elongation or differentiation of a bud, but once a tissue was fully developed, endophytes were no longer detected. Metabolic activity of the endophytes was suppressed at the onset of, and during, dormancy of Scots pine, but recovered before the following growing season.

In Situ Hybridization↗

Physiological responses of ponderosa pine in western Montana to thinning, prescribed fire and burning season.

Low-elevation ponderosa pine (Pinus ponderosa Dougl. ex. Laws.) forests of the northern Rocky Mountains historically experienced frequent low-intensity fires that maintained open uneven-aged stands. A century of fire exclusion has contributed to denser ponderosa pine forests with greater competition for resources, higher tree stress and greater risk of insect attack and stand-destroying fire. Active management intended to restore a semblance of the more sustainable historic stand structure and composition includes selective thinning and prescribed fire. However, little is known about the relative effects of these management practices on the physiological performance of ponderosa pine. We measured soil water and nitrogen availability, physiological performance and wood radial increment of second growth ponderosa pine trees at the Lick Creek Experimental Site in the Bitterroot National Forest, Montana, 8 and 9 years after the application of four treatments: thinning only; thinning followed by prescribed fire in the spring; thinning followed by prescribed fire in the fall; and untreated controls. Volumetric soil water content and resin capsule ammonium did not differ among treatments. Resin capsule nitrate in the control treatment was similar to that in all other treatments, although burned treatments had lower nitrate relative to the thinned-only treatment. Trees of similar size and canopy condition in the three thinned treatments (with and without fire) displayed higher leaf-area-based photosynthetic rate, stomatal conductance and mid-morning leaf water potential in June and July, and higher wood radial increment relative to trees in control units. Specific leaf area, mass-based leaf nitrogen content and carbon isotope discrimination did not vary among treatments. Our results suggest that, despite minimal differences in soil resource availability, trees in managed units where basal area was reduced had improved gas exchange and growth compared with trees in unmanaged units. Prescribed fire (either in the spring or in the fall) in addition to thinning, had no measurable effect on the mid-term physiological performance and wood growth of second growth ponderosa pine.

Fires↗

Effects of soil temperature and elevated atmospheric CO2 concentration on gas exchange, in vivo carboxylation and chlorophyll fluorescence in jack pine and white birch seedlings.

One-year-old jack pine (Pinus banksiana Lamb.) and current-year white birch (Betula papyrifera Marsh.) seedlings were grown in ambient (360 ppm) or twice ambient (720 ppm) atmospheric CO2 concentration ([CO2]) and at three soil temperatures (Tsoil = 7, 17 and 27 degrees C initially, increased to 10, 20 and 30 degrees C two months later, respectively) in a greenhouse for 4 months. In situ foliar gas exchange, in vivo carboxylation characteristics and chlorophyll fluorescence were measured after 2.5 and 4 months of treatment. Low Tsoil suppressed net photosynthetic rate (Pn), stomatal conductance (g(s)) and transpiration rate (E) in jack pine in both CO2 treatments and g(s) and E in white birch in ambient [CO2], but enhanced instantaneous water-use efficiency (IWUE) in both species after 2.5 months of treatment. Treatment effects on g(s) and E remained significant throughout the 4-month study. Low Tsoil reduced maximal carboxylation rate (Vcmax) and PAR-saturated electron transport rate (Jmax) in jack pine in elevated [CO2] after 2.5 months of treatment, but not after 4 months of treatment. Low Tsoil increased actual photochemical efficiency of photosystem II (PSII) in the light (DeltaF/Fm') in jack pine, but decreased DeltaF/Fm' in white birch after 4 months of treatment. In response to low Tsoil, photosynthetic linear electron transport to carboxylation (Jc) decreased in jack pine after 2.5 months and in white birch after 4 months of treatment. Low Tsoil increased the ratio of the photosynthetic linear electron transport to oxygenation (Jo) to the total photosynthetic linear electron transport rate through PSII (Jo/J(T)) in both species after 2.5 months of treatment, but the effects became statistically insignificant in white birch after 4 months of treatment. High Tsoil decreased foliar N concentration in white birch. Elevated [CO2] increased Pn, IWUE and Jc but decreased Jo/J(T) in both species at both measurement times except Jc in white birch after 2.5 months of treatment. Elevated [CO2] also decreased g(s) and E in white birch at high Tsoil, Vcmax in both species and triose phosphate utilization in white birch at low Tsoil after 4 months of treatment, and DeltaF/Fm' in white birch after 2.5 months of treatment. Elevated [CO2] also increased foliar N concentration in both species. Low Tsoil caused no permanent damage to PSII in either species, but jack pine responded and acclimated to low Tsoil more quickly than white birch. Photosynthetic down-regulation and a decrease in photosynthetic electron transport to photorespiration occurred in both species in response to elevated [CO2].

Betula↗