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Ozone uptake, water loss and carbon exchange dynamics in annually drought-stressed Pinus ponderosa forests: measured trends and parameters for uptake modeling.

This paper describes 3 years of physiological measurements on ponderosa pine (Pinus ponderosa Dougl. ex Laws.) growing along an ozone concentration gradient in the Sierra Nevada, California, including variables necessary to parameterize, validate and modify photosynthesis and stomatal conductance algorithms used to estimate ozone uptake. At all sites, gas exchange was under tight stomatal control during the growing season. Stomatal conductance was strongly correlated with leaf water potential (R2=0.82), which decreased over the growing season with decreasing soil water content (R2=0.60). Ozone uptake, carbon uptake, and transpirational water loss closely followed the dynamics of stomatal conductance. Peak ozone and CO2 uptake occurred in early summer and declined progressively thereafter. As a result, periods of maximum ozone uptake did not correspond to periods of peak ozone concentration, underscoring the inappropriateness of using current metrics based on concentration (e.g., SUM0, W126 and AOT40) for assessing ozone exposure risk to plants in this climate region. Both Jmax (maximum CO2-saturated photosynthetic rate, limited by electron transport) and Vcmax (maximum rate of Rubisco-limited carboxylation) increased toward the middle of the growing season, then decreased in September. Intrinsic water-use efficiency rose with increasing drought stress, as expected. The ratio of Jmax to Vcmax was similar to literature values of 2.0. Nighttime respiration followed a Q10 of 2.0, but was significantly higher at the high-ozone site. Respiration rates decreased by the end of the summer as a result of decreased metabolic activity and carbon stores.

California↗

Influence of climate on radial growth of Pinus cembra within the alpine timberline ecotone.

Radial growth variability and response to interannual climate variation of Cembran pine (Pinus cembra L.) were studied in the timberline ecotone on Mt. Patscherkofel (2246 m a.s.l.). The study area, which is in the inner alpine dry region of the Central Austrian Alps, is characterized by a continental climate with minimum precipitation in winter (about 150 mm during December-February) and frequent occurrence of warm dry winds (Föhn) in early spring. The hypothesis that spatial and temporal variability of radial growth is caused by site-related differences in sensitivity to winter stress (i.e., desiccation) was examined by applying dendroclimatological techniques. Ordination methods applied to tree ring time series revealed that spatial variability in radial growth is influenced by the local site factors elevation and slope aspect. Growth-climate relationships were explored using Pearson product-moment correlation coefficients and multiple regression analysis. Radial growth at the timberline was positively correlated with temperature in July and was also strongly correlated with mild temperatures in the previous autumn and high precipitation in winter (January-March). At the tree line, temperatures in the previous autumn and precipitation in late winter (March) also controlled radial growth, whereas July temperature was not significantly correlated with ring width. Because previous autumn temperature and winter precipitation were the main growth-determining factors at the timberline and the tree line, and both of these climate variables are known to influence susceptibility of trees to winter stress, the results support the working hypothesis. Analysis of climatic conditions in extreme growth years confirmed the high sensitivity of tree ring growth to precipitation in late winter (March) at the tree line plots. Furthermore, extent of growth reduction and release varied spatially and temporarily, with south- and west-facing stands showing a higher sensitivity to climate variation in the most recent decade (1990s) than the north-facing stand. This aspect-related change in sensitivity to climate may be associated with effects of climate warming on cambial activity.

Austria↗

Variations in dark respiration and mitochondrial numbers within needles of Pinus radiata grown in ambient or elevated CO2 partial pressure.

Within-leaf variations in cell size, mitochondrial numbers and dark respiration rates were compared in the most recently expanded tip, the mid-section and base of needles of Pinus radiata D. Don trees grown for 4 years in open-top chambers at ambient (36 Pa) or elevated (65 Pa) carbon dioxide partial pressure (p(CO2)a). Mitochondrial numbers and respiratory activity varied along the length of the needle, with the highest number of mitochondria per unit cytoplasm and the highest rate of respiration per unit leaf area at the base of the needle. Regardless of the location of the cells (tip, middle or basal sections), needles collected from trees grown in elevated p(CO2)a had nearly twice the number of mitochondria per unit cytoplasm as those grown in ambient p(CO2)a. This stimulation of mitochondrial density by growth at elevated p(CO2)a was greater at the tip of the needle (2.7 times more mitochondria than in needles grown in ambient CO2) than at the base of the needle (1.7 times). The mean size of individual mitochondria was unaffected either by growth at elevated p(CO2)a or by position along the needle. Tree growth at elevated p(CO2)a had a variable effect on respiration per unit leaf area, significantly increasing respiration in the tip of the needles (+25%) and decreasing respiration at the mid-section and base of the needles (-14% and -25%, respectively). Although a simple relationship between respiration per unit leaf area and mitochondrial number per unit cytoplasm was found within each CO2 treatment, the variable effect of growth at elevated p(CO2)a on respiration along the length of the needles indicates that a more complex relationship must determine the association between structure and function in these needles.

Carbon Dioxide↗

Seasonal variation of soil respiration in a Pinus cembra forest at the upper timberline in the Central Austrian Alps.

Soil respiration (R) of a 95-year-old Pinus cembra L. forest at the alpine timberline was measured continuously from October 2001 to January 2003 with an automated multiplexing gas exchange system. There was significant spatial variability in soil respiration, and R at a soil temperature of 10 degrees C (R10) decreased by about 20% m(-1) with increasing distance from the trunk. Needle litter and fine root density also decreased. The spatially averaged annual soil CO2 efflux was 35 g C m(-2) year(-1) in 2002. About 70% of the temporal variation in soil respiration could be explained by variations in soil temperature, whereas the influence of soil water potential and thus soil water content was negligible because soil water availability was supra-optimal.

Austria↗

Stomatal sensitivity to vapor pressure deficit and its relationship to hydraulic conductance in Pinus palustris.

We studied the response of stomatal conductance at leaf (gS) and canopy (GS) scales to increasing vapor pressure deficit (D) in mature Pinus palustris Mill. (longleaf pine) growing in a sandhill habitat in the coastal plain of the southeastern USA. Specifically, we determined if variation in the stomatal response to D was related to variation in hydraulic conductance along the soil-to-leaf pathway (KL) over the course of a growing season. Reductions in KL were associated with a severe growing season drought that significantly reduced soil water content (theta) in the upper 90-cm soil profile. Although KL recovered partially following the drought, it never reached pre-drought values. Stomatal sensitivity to D was well correlated with maximum gS at low D at both leaf and canopy scales, and KL appeared to influence this response by controlling maximum gS. Our results are consistent with the hypothesis that stomatal response to D occurs to regulate minimum leaf water potential, and that the sensitivity of this response is related to changes in whole-plant hydraulics.

Pinus↗

Spatial and seasonal variability of temperature responses of biochemical photosynthesis parameters and leaf nitrogen content within a Pinus densiflora crown.

We measured seasonal variation in area-based nitrogen concentration (N), maximum rate of carboxylation (Vcmax) and maximum rate of electron transport (Jmax) in 1-year-old needles along four first-order branches within a Pinus densiflora Sieb. & Zucc. crown, and analyzed their relationships to growth irradiance and temperature. Each leaf light environment was expressed as a ratio of the monthly mean of daily integrated photosynthetically active irradiance (Iint) for the particular needle to Iint above the canopy (Irel). Needle N decreased in the upper crown during the development of new needles, whereas it remained fairly constant in the lower crown, reflecting differences between upper and lower crown needles in their contribution to the nitrogen of new needles. Gradients of N within the crown were correlated with Irel in all seasons (r2 = 0.40-0.78). Seasonal variation in N was weakly correlated with mean daily air minimum temperatures. Both Vcmax and Jmax showed seasonal variation in all first-order branches, and decreased to their lowest values in winter. The gradients of Vcmax and Jmax within the crown were not correlated with Irel in some seasons, but were correlated with changes in N in most months (r2 = 0.33-0.75), except in the winter. Furthermore, the regression slope of the relationship between N and Vcmax and the temperature response of Vcmax and Jmax exhibited seasonal variation.

Electron Transport↗

Belowground carbon dynamics in loblolly pine (Pinus taeda) immediately following diammonium phosphate fertilization.

Forest soils store an immense quantity of labile carbon (C) and a may be a large potential sink for atmospheric C. Forest management practices such as fertilization may enhance overall C storage in soils, yet changes in physiological processes following nutrient amendments have not been widely investigated. We intensively monitored belowground C dynamics for nearly 200 days following diammonium phosphate fertilization of pot-grown loblolly pine (Pinus taeda L.) seedlings in an effort to examine the short-term effects of fertilization on processes involved in soil C sequestration. Soil respiration rates initially increased in fertilized pots relative to controls, followed by a brief reversal in this trend and then a final sustained pattern of elevated rates of soil respiration in the fertilized treatment. Patterns in soil respiration rates over time reflected changes in autotrophic (root) and heterotrophic (microbial) components of soil respiration. Root respiration rates were greater in the fertilized treatment 49 days following fertilization and returned to control rates by the end of the study. In contrast, microbial respiration rates and microbial activity per soil C concentration remained depressed over the same time period. Compared with control seedlings, total root biomass was 27% greater in fertilized seedlings harvested at the end of the study, indicating that the elevated soil respiration rates observed toward the end of the study were a result of increased respiring root biomass. We conclude that fertilization, at least over the short-term, may increase soil C sequestration by increasing belowground biomass production and reducing microbial driven C turnover.

Biomass↗

Canopy position affects photosynthetic adjustments to long-term elevated CO2 concentration (FACE) in aging needles in a mature Pinus taeda forest.

Few studies have examined the effects of elevated CO2 concentration ([CO2]) on the physiology of intact forest canopies, despite the need to understand how leaf-level responses can be aggregated to assess effects on whole-canopy functioning. We examined the long-term effects of elevated [CO2] (ambient + 200 ppm CO2) on two age classes of needles in the upper and lower canopy of Pinus taeda L. during the second through sixth year of exposure to elevated [CO2] in free-air (free-air CO2 enrichment (FACE)) in North Carolina, USA. Strong photosynthetic enhancement in response to elevated [CO2] (e.g., +60% across age classes and canopy locations) was observed across the years. This stimulation was 33% greater for current-year needles than for 1-year-old needles in the fifth and sixth years of treatment. Although photosynthetic stimulation in response to elevated [CO2] was maintained through the sixth year of exposure, we found evidence of concurrent down-regulation of Rubisco and electron transport capacity in the upper-canopy sunlit leaves. The lower canopy showed no evidence of down-regulation. The upper canopy down-regulated carboxylation capacity (Vcmax) and electron transport capacity (Jmax) by about 17-20% in 1-year-old needles; however, this response was significant across sampling years only for Jmax in 1-year-old needles (P < 0.02). A reduction in leaf photosynthetic capacity in aging conifer needles at the canopy top could have important consequences for canopy carbon balance and global carbon sinks because 1-year-old sunlit needles contribute a major proportion of the annual carbon balance of these conifers. Our finding of a significant interaction between canopy position and CO2 treatment on the biochemical capacity for CO2 assimilation suggests that it is important to take canopy position and needle aging into account because morphologically and physiologically distinct leaves could respond differently to elevated [CO2].

Carbon Dioxide↗

Organogenic responses of Pinus pinea cotyledons to hormonal treatments: BA metabolism and cytokinin content.

Isolated cotyledons from mature Pinus pinea L. embryos were cultured in vitro in a factorial combination of 4.4, 10 and 44.4 microM N6-benzyladenine (BA) for 2, 4, 8, 16 and 35 days to optimize shoot regeneration. Incubation of explants in 44.4 microM BA for 4 days, in place of the standard incubation in 4.4 microM BA for 35 days, reduced the entire culture period to 4 weeks. Shortening the culture period had no significant effect on the caulogenic response or the number of buds formed per cotyledon. To establish the relationship between key moments in the caulogenic process induced by 4.4 microM BA and the endogenous concentrations of the active forms of BA and other isoprenoid-type cytokinins (CKs), we examined uptake, metabolism and amount of BA, as well as the amounts of zeatin, dihydrozeatin and their ribosides in P. pinea cotyledons after 1, 2, 6, 12 and 24 h, and 2, 4, 8, 16 and 35 days of exposure to 8-[14C]BA. Uptake and release of BA were associated with water movement between explants and the medium during the first 8 days of culture. The interconvertible forms of BA were the main metabolites formed in the tissues. Inactivation of BA as a result of conjugation or oxidation was insignificant. The endogenous concentration of BA + N6-benzyladenosine was 20-fold higher than the exogenously applied BA during the competence acquisition phase (Days 0-3). The concentration of isoprenoid-type CKs also increased 16-fold and then decreased during this time. Induction of shoot buds (Days 4-8) was characterized by a second peak of BA uptake by explants that triggered the synthesis of N6-benzyladenosine-5 -monophosphate and by the maintenance of isoprenoid-type CKs. Reestablishment of CK homeostasis marked the shift from the induction phase to the shoot development phase in this organogenic process (Days 8-12).

Benzyl Compounds↗

Somatic embryogenesis from vegetative shoot apices of mature trees of Pinus patula.

Embryogenic cultures were initiated and established from apical shoots of mature trees of three genotypes of Pinus patula Scheide et Deppe. Factors affecting initiation, including cold pretreatment, basal medium composition, growth regulators and gelling agent concentration, and the effect of partial desiccation on somatic embryo maturation were investigated. Cold pretreatment of thick sections (0.5-1.0 mm) of apical shoots at 2 degrees C for 3 days on 0.3% activated charcoal induced white mucilaginous embryogenic callus on initiation medium. Subculture of this embryogenic callus on maintenance medium resulted in the formation of embryonal suspensor masses with proembryos. Partial desiccation (12-90 h) of embryogenic tissue at the proembryo stage of development, prior to transfer to maturation medium containing 9 g l(-1) Gellan gum, enhanced somatic embryo maturation and germinability. The frequency of maturation increased from 5.3 to 16.5% after 12 h of desiccation and from 16.5 to 73.8% after 24 h of desiccation, but longer periods of desiccation were ineffective.

Pinus↗

Variation in needle longevity is related to needle-fascicle production rate in Pinus sylvestris.

Latitudinal variation in needle longevity of conifers in response to climatic variability is a well-known phenomenon, but its significance has only rarely been studied. Scots pine (Pinus sylvestris L.) stands were investigated in four locations in Lapland (northern Finland and northern Sweden, 67-68 degrees N) and in four locations in Estonia (59 degrees N) to test the hypothesis that plasticity in needle longevity results in similar needle biomass per foliated shoot length across latitudes. The needle-trace method (NTM) revealed the dynamics of needle fascicles attached to stem shoots. Long-term mean needle age was 2 years greater in Lapland than in Estonia. In both regions, mean needle age was strongly and negatively related to yearly needle-fascicle production rate relative to the number of needles on the same shoot axis. Although significantly fewer needle-fascicles were produced annually in trees in Lapland than in Estonia, the overall number of fascicles attached to the stem shoots, needle-fascicle density and individual needle dry mass did not differ between regions. Consequently, needle biomass per foliated shoot length was similar in trees in both regions. Thus, our results support the theory that plasticity in needle longevity helps Scots pine to compensate for reduced needle production so that the same foliar biomass per shoot is retained under a wide range of growth conditions.

Climate↗

Carbon budget of Pinus sylvestris saplings after four years of exposure to elevated atmospheric carbon dioxide concentration.

To study the responses of Scots pine (Pinus sylvestris L.), a commercially important tree species in Europe, to future increases in atmospheric CO2 concentration ([CO2]), we grew saplings for 4 years in the ground in open-top chambers in ambient or ambient + 400 micromol mol(-1) CO2, without supplemental addition of nutrients and water. Carbon (C) budgets were developed for trees in both CO2 treatments based on productivity and biomass data obtained from destructive harvests at the end of the third and fourth years of treatment, and simulations of annual gross photosynthesis (P(tot)) and maintenance respiration by the model MAESTRA. Simulated P(tot) was enhanced by elevated [CO2], despite significant down-regulation of photosynthetic capacity. The subsequent increase in C uptake was allocated primarily to tissues with limited longevity (needles and fine roots), which explains why the measured annual increment in woody biomass did not differ between CO2 treatments. Thus, our results suggest that accelerated stem growth only occurs in the first 2 years in the presence of elevated [CO2] and that forest rotations will not be shortened significantly in response to increasing [CO2]. In elevated [CO2], a higher proportion of available C was allocated below ground, resulting in altered biomass distribution patterns. In trees of equal size, measured ratios of fine root/needle biomass and belowground/aboveground biomass were almost twice as large in the elevated [CO2] treatment. Although there are uncertainties in scaling from saplings to mature canopies, the data indicate that, in nutrient-limited Scots pine forests, elevated [CO2] is unlikely to accelerate tree growth significantly, but is likely to increase C inputs to soil.

Biomass↗

Contrasting net primary productivity and carbon distribution between neighboring stands of Quercus robur and Pinus sylvestris.

Standing biomass, net primary production (NPP) and soil carbon (C) pools were studied in a 67-year-old pedunculate oak (Quercus robur L.) stand and a neighboring 74-year- old Scots pine (Pinus sylvestris L.) stand in the Belgian Campine region. Despite a 14% lower tree density and a lower tree height in the oak stand, standing biomass was slightly higher than in the pine stand (177 and 169 Mg ha(-1) in oaks and pines, respectively), indicating that individual oak trees contained more biomass than pine trees of similar diameter. Moreover, NPP in the oak stand was more than double that in the pine stand (17.7 and 8.1 Mg ha(-1) year(-1), respectively). Several observations indicated that soil organic matter accumulated at higher rates under pines than under oaks. We therefore hypothesized that the pines were exhibiting an age-related decline in productivity due to nutrient limitation. The poor decomposability of pine litter resulted in the observed accumulation of organic matter. The subsequent immobilization of nutrients in the organic matter, combined with the already nutrient-poor soil conditions, resulted in a decrease in total NPP over time, as well as in a substantial shift in the allocation of NPP toward fine roots. In the oak stand, litter is less recalcitrant to decay and soil acidity is less severe; hence, organic matter does not accumulate and nutrients are recycled. This probably explains why NPP was much higher in the oaks than in the pines and why only a small proportion of NPP was allocated to oak fine roots.

Biomass↗

Analysis of biomass accumulation and stem size distributions over long periods in managed stands of Pinus sylvestris in Finland using the 3-PG model.

We tested the performance of a process-based model (PBM) in relation to long-term mensuration data from two sites in Finland where the stands were up to 90 years old and had been thinned at approximately 5-year intervals over the last 50 years. The PBM used was based on the 3-PG (Physiological Principles to Predict Growth) model developed by Landsberg and Waring (1997), with modifications in the biomass allocation routine, for which we used data and calculations by Vanninen (2003) to estimate the allocation coefficients and turnover rates. Site fertility was estimated in terms of known site-type characteristics. The model was evaluated in terms of stand development and its ability to simulate responses to thinning; stem numbers after thinning were specified at the dates when the thinning took place. Stand development in terms of basal area, volume and mean diameter at breast height, closely followed the measured characteristics of all stands. Foliage mass predictions were close to estimates obtained by an empirical method. The analysis shows that, under normal thinning regimes, a range of different thinning intensities can be adequately described using a simple multiplicative model relating the proportion of volume and foliage mass removed to the corresponding proportion of stem numbers. This model, together with stem allometry data, described the "growth" in mean diameter after thinning, which simply reflected the removal of the smaller trees. These results indicate that, with a single set of parameter values, 3-PG can provide good descriptions of the growth patterns of trees-in this case Pinus sylvestris L.-over long periods, including growth after repeated thinning. One of the outputs from the 3-PG model is mean stem diameter (B): we show that it is feasible to estimate stem size distributions, which changed considerably over the life of these stands, from B using the Weibull function. This shows that, given information about the Weibull parameters for particular species and cultural systems, it should be possible to use stem numbers and the B obtained from the 3-PG model to produce information about stem size distributions from simulated data.

Biomass↗

Estimating foliage biomass in Scots pine (Pinus sylvestris) and Norway spruce (Picea abies) plots.

Dynamic decomposition models are needed to estimate changes in the carbon stock of boreal soil because these changes are difficult to measure directly. An important aboveground carbon flux to the soil is foliage litterfall. To estimate this flux, both the amount and the turnover rate of the foliage biomass component must be known. Several methods for estimating foliage biomass of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.), including biomass equations and biomass expansion factors (BEFs), were compared with predicted foliage biomass based on forest inventory plot-level measurements. Measured foliage biomass was up-scaled from the branch-level to the plot-level by combining forest inventory variables (diameter, height, height at the crown base and crown base diameter) based on the assumptions of pipe model theory. Combining the foliage biomass: cross-sectional area ratio with the forest inventory variables provided accurate estimates of foliage biomass at the plot-level for plots in southern Finland. The results emphasize the need to test biomass equations with independent data, especially when the equations applied are based on neighboring regions.

Biomass↗

Seasonal variation in gene expression for loblolly pines (Pinus taeda) from different geographical regions.

In developing xylem, gene expression levels vary in different genotypes, at different stages of development, throughout a growing season, and in response to stresses. Commercially important characteristics such as wood-specific gravity are known to differ with seed source. For example, when grown on a common site, the specific gravity of Arkansas loblolly pine (Pinus taeda L.) trees is greater than that of Louisiana loblolly pine, and Texas loblolly pines have a greater specific gravity than loblolly pines from the Atlantic coast. A microarray analysis was performed to examine variation in gene expression among trees from different geographical sources when grown on a common site, and seasonal variation in gene expression in each seed source. We used microarrays containing 2171 expressed sequence tags (ESTs) with putative functions of interest, selected from several loblolly pine xylem partial cDNA libraries and a shoot tip library. Genes with significant variation in expression for each factor were identified. Many genes preferentially expressed in latewood compared with earlywood were for proteins involved in cell wall biosynthesis. Variation in gene expression among trees from the two seed sources in each growing season suggests that there may be more differences between South Arkansas trees and South Louisiana trees in latewood than in earlywood. Variation in gene expression among trees from different regions may reflect adaptation to different environments.

Gene Expression Profiling↗

Elevated atmospheric CO2 concentration alters the effect of phosphate supply on growth of Japanese red pine (Pinus densiflora) seedlings.

We demonstrated that the inorganic phosphate (P(i)) requirement for growth of Japanese red pine (Pinus densiflora Sieb. & Zucc.) seedlings is increased by elevated CO(2) concentration ([CO(2)]) and that responses of the ectomycorrhizal fungus Pisolithus tinctorius (Pers.) Coker & Couch to P(i) supply are also altered. To investigate the growth response of non-mycorrhizal seedlings to P(i) supply in elevated [CO(2)], non-mycorrhizal seedlings were grown for 73 days in ambient or elevated [CO(2)] (350 or 700 micromol mol(-1)) with nutrient solutions containing one of seven phosphate concentrations (0, 0.02, 0.04, 0.06, 0.08, 0.10 and 0.20 mM). In ambient [CO(2)], the growth response to P(i) was saturated at about 0.1 mM P(i), whereas in elevated [CO(2)], the growth response to P(i) supply did not saturate, even at the highest P(i) supply (0.2 mM), indicating that the P(i) requirement is higher in elevated [CO(2)] than in ambient [CO(2)]. The increased requirement was due mainly to an altered shoot growth response to P(i) supply. The enhanced P(i) requirement in elevated [CO(2)] was not associated with a change in photosynthetic response to P(i) or a change in leaf phosphorus (P) status. We investigated the effect of P(i) supply (0.04, 0.08 and 0.20 mM) on the ectomycorrhizal fungus P. tinctorius in mycorrhizal seedlings grown in ambient or elevated [CO(2)]. Root ergosterol concentration (an indicator of fungal biomass) decreased with increasing P(i) supply in ambient [CO(2)], but the decrease was far less in elevated [CO(2)]. In ambient [CO(2)] the ratio of extramatrical mycelium to root biomass decreased with increasing P(i) supply but did not change in elevated [CO(2)]. We conclude that, because elevated [CO(2)] increased the P(i) requirement for shoot growth, the significance of the ectomycorrhizal association was also increased in elevated [CO(2)].

Atmosphere↗

Righting response of artificially inclined maritime pine (Pinus pinaster) saplings to wind loading.

To determine if trees respond to dynamic and static loading in the same manner, 2-year-old maritime pine (Pinus pinaster Ait.) trees were subjected to different types of mechanical loading in the field. One block of trees (the control) were kept in pots and planted in the field at an angle of 0 or 45 degrees to the vertical. A similar block of leaning potted trees was planted nearby and subjected to frequent, unilateral wind loading for a period of 1 s every 2 min. Half the leaning trees were oriented toward the direction of wind loading and half were oriented along the axis of wind loading. The stem profile was measured three times during the growing season to quantify the rate of stem straightening. Compression wood formation and stem shape were measured in all plants. No differences in mean height or diameter were observed between blocks and all leaning trees straightened, but not at the same rate. Although no difference in the rate of apical straightening occurred between control and wind-treated trees, the righting response of the basal part of the stem of leaning trees subjected to wind was four times greater than that of leaning trees without wind. No differences in the righting response were observed between leaning trees growing toward and trees growing away from the source of wind. No significant differences in compression wood formation were found between control trees and wind-treated trees, indicating that other factors must determine the reorientation rate of leaning trees. Results are discussed with reference to the quality of compression wood in conifers and the mechanotransductive pathway in plants.

Pinus↗