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PubMed2004

Seasonal and annual stem respiration of Scots pine trees under boreal conditions.

BACKGROUND AND AIMS: Stem respiration of trees is a major, but poorly assessed component of the carbon balance of forests, and important for geo-chemistry. Measurements are required under naturally changing seasonal conditions in different years. Therefore, intra- and inter-annual carbon fluxes of stems in forests were measured continuously from April to November in three consecutive years. METHODS: Stem respiratory CO2 fluxes of 50-year-old Scots pine (Pinus sylvestris) trees were continuously measured with a CO2 analyser, and, concomitantly, stem circumference, stem and air temperature and other environmental factors and photosynthesis, were also measured automatically. KEY RESULTS: There were diurnal, seasonal and inter-annual changes in stem respiration, which peaked at 1600 h during the day and was highest in July. The temperature coefficient of stem respiration (Q10) was greater during the growing season than when growth was slow or had stopped, and more sensitive to temperature in the growing season. The annual Q10 remained relatively constant at about 2 over the three years, while respiration at a reference temperature of 15 degrees C (R15) was higher in the growing than in the non-growing season (1.09 compared with 0.78 micromol m(-2) stem surface s(-1)), but was similar between the years. Maintenance respiration was 76 %, 82 % and 80 % of the total respiration of 17.46, 17.26 and 19.35 mol m2 stem surface in 2001, 2002 and 2003, respectively. The annual total stem respiration of the stand per unit ground area was 75.97 gC m(-2) in 2001 and 74.28 gC m(-2) in 2002. CONCLUSIONS: Stem respiration is an important component in the annual carbon balance of a Scots pine stand, contributing 9 % to total carbon loss from the ecosystem and consuming about 8 % of the carbon of the ecosystem gross primary production. Stem (or air) temperature was the most important predictor of stem carbon flux. The magnitude of stem respiration is modified by photosynthesis and tree growth. Solar radiation indirectly affects stem respiration through its effect on photosynthesis.

Carbon↗

PubMed2007

Total and component carbon fluxes of a Scots pine ecosystem from chamber measurements and eddy covariance.

BACKGROUND AND AIMS: Distinguishing between, and quantifying, the different components of ecosystem C fluxes is critical in predicting the responses of ecosystem C cycling to climate change. The aims of this study were to quantify the photosynthetic and respiratory fluxes of a 50-year-old Scots pine (Pinus sylvestris) ecosystem, and to distinguish respiration of branches with needles from that of stems, and that of soil. METHODS: The CO2 flux of the ecosystem was continuously measured using the eddy covariance (EC) method, and its components (respiration and photosynthesis of a branch with needles, stem and soil surface) were measured with an automated chamber system, from 2001 to 2004. KEY RESULTS: All values below are chamber based. The average temperature coefficient (Q10) of respiration was 2.7, 2.2 and 4.0, respectively, for branch (Rbran), stem (Rstem) and the soil surface (Rsoil). Respiration at a reference temperature of 15 degrees C (R15) was 1.27, 0.49 and 4.02 micromol CO2 m(-2) ground s(-1) for the three components, respectively. Over 4 years, the annual Rbran, Rstem and Rsoil ranged from 196 to 256, 56 to 83 and 439 to 598 g C m(-2) ground year(-1), respectively, with a 4-year average of 227, 72 and 507 g C m(-2) ground year(-1). Annual ecosystem respiration (Reco) was 731, 783, 909 and 751 g C m(-2) ground year(-1) in years 2001-2004, respectively, gross primary production (GPP) was 922, 1030, 1138 and 1001 g C m(-2) ground year(-1), and net ecosystem production (NEP) was 191, 247, 229 and 251 g C m(-2) ground year(-1). The average contribution of Rbran, Rstem and Rsoil to Reco was 29, 9 and 62 %, respectively. Overstorey photosynthesis accounted for 96 % of GPP. The average Reco/GPP ratio was 0.78. Net primary production (NPP) in the 4 years was 469, 581, 600 and 551 g C m(-2) year(-1), respectively, with the NPP/GPP ratio 0.54 averaged over the years. CONCLUSIONS: Respiration from the soil is the dominant component of ecosystem respiration. Differences between years in Reco were due to differences in temperature during the growing season. Rsoil was more sensitive to temperature than Rbran and Rstem, and differences in Rsoil were responsible for the differences in Reco between years.

Carbon↗

PubMed1994

Identification of quantitative trait loci influencing wood specific gravity in an outbred pedigree of loblolly pine.

We report the identification of quantitative trait loci (QTL) influencing wood specific gravity (WSG) in an outbred pedigree of loblolly pine (Pinus taeda L.). QTL mapping in an outcrossing species is complicated by the presence of multiple alleles (> 2) at QTL and marker loci. Multiple alleles at QTL allow the examination of interaction among alleles at QTL (deviation from additive gene action). Restriction fragment length polymorphism (RFLP) marker genotypes and wood specific gravity phenotypes were determined for 177 progeny. Two RFLP linkage maps were constructed, representing maternal and paternal parent gamete segregations as inferred from diploid progeny RFLP genotypes. RFLP loci segregating for multiple alleles were vital for aligning the two maps. Each RFLP locus was assayed for cosegregation with WSG QTL using analysis of variance (ANOVA). Five regions of the genome contained one or more RFLP loci showing differences in mean WSG at or below the P = 0.05 level for progeny as grouped by RFLP genotype. One region contained a marker locus (S6a) whose QTL-associated effects were highly significant (P > 0.0002). Marker S6a segregated for multiple alleles, a prerequisite for determining the number of alleles segregating at the linked QTL and analyzing the interactions among QTL alleles. The QTL associated with marker S6a appeared to be segregating for multiple alleles which interacted with each other and with environments. No evidence for digenic epistasis was found among the five QTL.

Alleles↗

PubMed2000

Whole-genome characterization of embryonic stage inbreeding depression in a selfed loblolly pine family.

Inbreeding depression is important in the evolution of plant populations and mating systems. Previous studies have suggested that early-acting inbreeding depression in plants is primarily due to lethal alleles and possibly epistatic interactions. Recent advances in molecular markers now make genetic mapping a powerful tool to study the genetic architecture of inbreeding depression. We describe a genome-wide evaluation of embryonic viability loci in a selfed family of loblolly pine (Pinus taeda L.), using data from AFLP markers from an essentially complete genome map. Locus positions and effects were estimated from segregation ratios using a maximum-likelihood interval mapping procedure. We identified 19 loci showing moderately deleterious to lethal embryonic effects. These loci account for >13 lethal equivalents, greater than the average of 8.5 lethal equivalents reported for loblolly pine. Viability alleles show predominantly recessive action, although potential overdominance occurs at 3 loci. We found no evidence for epistasis in the distribution of pairwise marker correlations or in the regression of fitness on the number of markers linked to deleterious alleles. The predominant role of semilethal alleles in embryonic inbreeding depression has implications for the evolution of isolated populations and for genetic conservation and breeding programs in conifers.

Crosses, Genetic↗

PubMed2001

Cytonuclear disequilibrium and genetic drift in a natural population of ponderosa pine.

We measured the cytonuclear disequilibrium between 11 nuclear allozyme loci and both mitochondrial and chloroplast DNA haplotypes in a natural population of ponderosa pine (Pinus ponderosa, Laws). Three allozyme loci showed significant associations with mtDNA variation, while two other loci showed significant association with cpDNA. However, the absolute number of individuals involved in any of the associations was small, such that in none of the nuclear-organellar combinations was the difference between observed and expected numbers >11 individuals. Patterns of association were not consistent across loci or organellar genomes, suggesting that they are not the result of mating patterns, which would act uniformly on all loci. This pattern of disequilibria is consistent with the action of genetic drift and with existing knowledge of the structure of this population and thus does not imply the action of other evolutionary processes. The overall magnitude (normalized disequilibrium) of associations was greater for maternally inherited mtDNA than for paternally inherited cpDNA, though this difference was neither large nor significant. Such significant disequilibria involving the paternally inherited organelle indicate that not only are there a limited number of seed parents, but the effective number of pollen parents is also limited.

Cell Nucleus↗

PubMed2001

Bayesian methods for quantitative trait loci mapping based on model selection: approximate analysis using the Bayesian information criterion.

We describe an approximate method for the analysis of quantitative trait loci (QTL) based on model selection from multiple regression models with trait values regressed on marker genotypes, using a modification of the easily calculated Bayesian information criterion to estimate the posterior probability of models with various subsets of markers as variables. The BIC-delta criterion, with the parameter delta increasing the penalty for additional variables in a model, is further modified to incorporate prior information, and missing values are handled by multiple imputation. Marginal probabilities for model sizes are calculated, and the posterior probability of nonzero model size is interpreted as the posterior probability of existence of a QTL linked to one or more markers. The method is demonstrated on analysis of associations between wood density and markers on two linkage groups in Pinus radiata. Selection bias, which is the bias that results from using the same data to both select the variables in a model and estimate the coefficients, is shown to be a problem for commonly used non-Bayesian methods for QTL mapping, which do not average over alternative possible models that are consistent with the data.

Alleles↗

PubMed2004

A new method for loading Bursaphelenchus xylophilus (Nematoda: Aphelenchoididae) on adult Monochamus alternatus (Coleoptera: Cerambycidae).

A new method was developed for loading the pinewood nematode, Bursaphelenchus xylophilus (Steiner et Buhrer) Nickle, on the beetle Monochamus alternatus Hope. Postdiapause beetle larvae were sterilized with 70 and 99.9% aqueous ethanol and placed singly in flasks where B. xylophilus reproduced on the fungus Ophiostoma minus (Hedgcock) H. et P. Sydow that had been grown on autoclaved barley grain and Pinus densifiora Sieb. et Zucc. wood chips. The fungus produced a large nematode population that developed to a high proportion of third-stage dispersal juveniles that molted to the fourth-stage dispersal juveniles. The survival rate was 80%, and the mean nematode load was 10,096. It took a mean of 5 wk to obtain the nematode-infested beetles after the initiation of nematode rearing.

Animals↗

PubMed2004

Long-term benefits to the growth of ponderosa pines from controlling southwestern pine tip moth (Lepidoptera: Tortricidae) and weeds.

The southwestern pine tip moth, Rhyacionia neomexicana (Dyar) (Lepidoptera: Tortricidae), is a native forest pest that attacks seedlings and saplings of ponderosa pine, Pinus ponderosa Dougl. ex Laws, in the southwestern United States. Repeated attacks can cause severe deformation of host trees and significant long-term growth loss. Alternatively, effective control of R. neomexicana, vegetative competition, or both in young pine plantations may increase survival and growth of trees for many years after treatments are applied. We test the null hypothesis that 4 yr of R. neomexicana and weed control with insecticide, weeding, and insecticide plus weeding would not have any residual effect on survival and growth of trees in ponderosa pine plantation in northern Arizona 14 yr post-treatment, when the trees were 18 yr old. Both insecticide and weeding treatment increased tree growth and reduced the incidence of southwestern pine tip moth damage compared with the control. However, weeding alone also significantly increased tree survival, whereas insecticide alone did not. The insecticide plus weeding treatment had the greatest tree growth and survival, and the lowest rate of tip moth damage. Based on these results, we rejected our null hypothesis and concluded that there were detectable increases in the survival and growth of ponderosa pines 14 yr after treatments applied to control R. neomexicana and weeds.

Animals↗

PubMed2005

Evaluation of biological and biorational control tactics for suppression of Nantucket pine tip moth damage in Virginia pine Christmas trees.

A study was performed to evaluate the potential of the egg parasitoid Trichogramma exiguum Pinto and Platner (Hymenoptera: Trichogrammatidae), the newly registered insect growth regulator (IGR) tebufenozide (Confirm), and a modified spray technique (top whorl only pesticide application) for suppression of Nantucket pine tip moth, Rhyacionia frustrana (Comstock) (Lepidoptera: Tortricidae), damage in Virginia pine, Pinus virginiana Mill., Christmas trees. Augmentative releases of T. exiguum failed to increase parasitism levels in release plots compared with controls, and significant reduction in tip moth damage did not occur. High predation levels on released T. exiguum may have contributed to the failure of parasitoid augmentations. Whole-tree and top whorl tebufenozide treatments provided significantly greater damage control than corresponding applications of acephate (Orthene), a commonly used pesticide in Christmas trees. Damage to trees receiving whole-tree chemical applications did not differ significantly from trees receiving top whorl treatments for most measurements of damage. Top whorl chemical treatments resulted in a 67% reduction in time required for application and a 70% reduction in pesticide used.

Animals↗

PubMed2005

Phenology, natural enemies, and efficacy of horticultural oil for control of Chionaspis heterophyllae (Homoptera: Diaspididae) on Christmas tree plantations.

Pine needle scale, Chionaspis pinifoliae (Fitch), and Chionaspis heterophyllae Cooley are important pests of Scots pine, Pinus sylvestris L., and other conifers in much of North America. On Christmas tree plantations, these insects are typically controlled by spraying broad-spectrum insecticides when the vulnerable immature stages are present. However, effective control of bivoltine populations can be difficult to achieve due to asynchronous hatch and development of the second generation. Our objectives were to 1) determine the phenology of the second generation of C. heterophyllae in Michigan; 2) characterize the natural enemy complex; and 3) assess the effectiveness of horticultural oil for control of C. heterophyllae on P. sylvestris Christmas tree plantations. We monitored scale populations in three counties in lower Michigan for 3 yr. Scale phenology was consistently associated with cumulative degree-days base 10 degrees C (DD(10 degrees C)). Second-generation egg hatch began at approximately 1230-1300 DD(10 degrees C), and continued for approximately 3 wk. The peak of the second instar coincided with 1500-1600 DD(10 degrees C). Common predators included the coccinellids Chilocorus stigma (Say) and Microweisia misella (LeConte). On average, 70% of the C. heterophyllae population in unsprayed fields was killed by predators in 1999. Two endoparasitic wasps, Encarsia bella Gahan and Marietta mexicana Howard (Hymenoptera: Aphelinidae), also were recovered. In 2000 and 2001, we applied a highly refined horticultural spray oil with a backpack mist blower at 1500-1600 DD(10 degrees). Scale mortality on trees treated with oil ranged from 66 to 80% and was similar to control achieved using conventional insecticides in both years.

Animals↗

PubMed2006

Efficacy of systemic insecticides for protection of loblolly pine against southern pine engraver beetles (Coleoptera: Curculionidae: Scolytinae) and wood borers (Coleoptera: Cerambycidae).

We evaluated the efficacy of the systemic insecticides dinotefuran, emamectin benzoate, fipronil, and imidacloprid for preventing attacks and brood production of southern pine engraver beetles (Coleoptera: Curculionidae: Scolytinae) and wood borers (Coleoptera: Cerambycidae) on standing, stressed trees and bolt sections of loblolly pine, Pinus taeda L., in eastern Texas. Emamectin benzoate significantly reduced the colonization success of engraver beetles and associated wood borers in both stressed trees and pine bolt sections. Fipronil was nearly as effective as emamectin benzoate in reducing insect colonization of bolts 3 and 5 mo after injection but only moderately effective 1 mo after injection. Fipronil also significantly reduced bark beetle-caused mortality of stressed trees. Imidacloprid and dinotefuran were ineffective in preventing bark beetle and wood borer colonization of bolts or standing, stressed trees. The injected formulation of emamectin benzoate was found to cause long vertical lesions in the sapwood-phloem interface at each injection point.

Animals↗

PubMed2001

Possible explanation of the disparity between the in vitro and in vivo measurements of Rubisco activity: a study in loblolly pine grown in elevated pCO2.

Rubisco activity can be measured using gas exchange (in vivo) or using in vitro methods. Commonly in vitro methods yield activities that are less than those obtained in vivo. Rubisco activity was measured both in vivo and in vitro using a spectrophotometric technique in mature Pinus taeda L. (loblolly pine) trees grown using free-air CO2 enrichment in elevated (56 Pa) and current (36 Pa) pCO2. In addition, for studies where both in vivo and in vitro values of Rubisco activity were reported net CO2 uptake rate (A) was modelled based on the in vivo and in vitro values of Rubisco activity reported in the literature. Both the modelling exercise and the experimental data showed that the in vitro values of Rubisco activity were insufficient to account for the observed values of A. A trichloroacetic acid (TCA) precipitation of the protein from samples taken in parallel with those used for activity analysis was co-electrophoresed with the extract used for determining in vitro Rubisco activity. There was significantly more Rubisco present in the TCA precipitated samples, suggesting that the underestimation of Rubisco activity in vitro was attributable to an insufficient extraction of Rubisco protein prior to activity analysis. Correction of in vitro values to account for the under-represented Rubisco yielded mechanistically valid values for Rubisco activity. However, despite the low absolute values for Rubisco activity determined in vitro, the trends reported with CO2 treatment concurred with, and were of equal magnitude to, those observed in Rubisco activity measured in vivo.

Carbon Dioxide↗

PubMedDate not supplied

On the number of genes controlling the grass stage in longleaf pine.

The grass stage is an inherent and distinctive developmental trait of longleaf pine (Pinus palustris), in which height growth in the first few years after germination is suppressed. In operational forestry practice the grass stage extends for two to several years and often plays a role in planting failures and decisions to plant alternative species. Interspecies hybrids involving loblolly (P. taeda) and slash (P. elliottii var. elliottii) pines have been investigated as a means to produce planting stock with improved early height growth and to develop backcross populations for advanced generation breeding. We have reevaluated data from several interspecies populations, with the objective of estimating the number of genes contributing to the difference in first-year height growth between longleaf and loblolly pines. Estimates based on means and variances of parental and interspecies hybrid and backcross families suggest a minimum of 4 to 10 genes with standard errors less than half the estimates. These results suggest that the grass stage has evolved through the accumulation of alleles at several loci, each with small effects on various components of first-year height growth. Given the complexity of the grass-stage trait, tree breeders may need to combine genetic marker analysis with recurrent backcross breeding to efficiently develop longleaf pine planting stock for improved reforestation.

Analysis of Variance↗

PubMed2005

Influence of meso- and microscale habitat structure on focal distribution of sympatric Ixodes scapularis and Amblyomma americanum (Acari: Ixodidae).

We compared the distribution of sympatric Ixodes scapularis Say and Amblyomma americanum (L.) within several suitable forested habitats at different spatial scales and characterized differences in microhabitat features accounting for the observed distribution of questing ticks. We used automatic data loggers placed in the shrub and litter layers to contrast mesoclimate and microclimate conditions experienced by questing ticks. Larger numbers of I. scapularis were collected at sites where the forest canopy was more fragmented and where the canopy contained more hardwood species than pitch pine, Pinus rigida Mill. Dominance of pine in the canopy affected the character of the shrub layer vegetation and composition of the litter layer, which concomitantly affected the microclimate conditions experienced by questing ticks. Pitch pine-dominated habitats were drier and hotter than those under a broad-leaved canopy, and questing ticks experienced increased saturation deficit in the later spring and summer in pine forest sites. The shrub layer vegetation seemed to have a moderating effect on the microclimate experienced by questing ticks and subtle differences in vegetation structure resulted in substantially different conditions as encountered by questing ticks over the space of a few meters. In contrast to questing I. scapularis, all three stages of questing A. americanum exhibited poor relationships with microclimate variables recorded in the litter and shrub layers. Further research is required to determine which environmental conditions and which habitats are most likely to support this species.

Animals↗

PubMed2002

The fungal sheath of ectomycorrhizal pine roots: an apoplastic barrier for the entry of calcium, magnesium, and potassium into the root cortex?

The apoplastic permeability of the fungal sheath of two different ectomycorrhizal associations of Pinus sylvestris L. was analysed by laser microprobe mass analysis (LAMMA) and energy-dispersive X-ray spectroscopy (EDXS) after stable isotope labelling with 25Mg, 41K and 44Ca. Entry of 25Mg and 44Ca into the outer cortical apoplast of non-mycorrhizal roots was detected after 4 min of labelling. After a longer exposure time the endodermis with its Casparian band acted as an efficient apoplastic diffusion barrier for the radial movement of 25Mg and 44Ca into the stele. A fraction of approximately one-third of the apoplastic cations of the root cortex could not be exchanged against the external label even after longer exposure times. The ectomycorrhizal sheath of the two fungal species used, Pisolithus tinctorius (Pers.) Coker & Couch and Suillus bovinus (L. ex Fr.) Kuntze, does not completely inhibit the apoplastic movement of ions into the mycorrhizal root cortex, but retarded the penetration of isotopes into the cortical apoplast. In roots inoculated with S. bovinus, a clear labelling of the cortical apoplast could first be detected after 24 h of exposure to the stable isotope solution. At this time the labelling of the cortical apoplast in these mycorrhizal roots was higher than those of non-mycorrhizal roots and, with EDXS, changes in the element composition of the apoplast were detected. The results indicated that possibly hydrophobins localized in the fungal cell wall might be involved in the increased hydrophobicity of mycorrhizal roots and the lower permeability of the ectomycorrhizal sheath.

Calcium↗

PubMed2004

Annual and seasonal variation of sap flow and conductance of pine trees grown in elevated carbon dioxide and temperature.

Measurements of sap flow, crown structure, and microclimate were used to estimate the transpiration of individual 30-year-old Pinus sylvestris L. trees grown in elevated temperature and CO2. The trees were enclosed in closed-top chambers and exposed either to current ambient conditions (CON), or elevated CO2 (+350 micromol mol(-1); EC), or elevated temperature (+2 to +6 degrees C; ET) or a combination of EC and ET (ECT) since 1996, and the measurements were made from 1999 to 2001. EC significantly increased annual sap flow per tree (Ft.m) by 14% in 1999, but reduced it by 13% in 2000 and 16% in 2001. The CO2-induced increase in Ft.m in 1999 was due to a large increase in foliage area of trees, which more than compensated for a small decrease in crown conductance (Gc). The CO2-induced decreases in Ft.m in 2000 and 2001 resulted from a pronounced decline in Gc, which was much greater than the increase in foliage area. The CO2-induced increase in sensitivity of Gc at high vapour pressure deficit (VPD) did not alter the general response of sap flow to CO2 enrichment, but it did affect the diurnal courses of sap flow on some days during the main growing season (days 150-240). ET increased Ft.m by 53%, 45%, and 57% in 1999, 2000, and 2001, respectively, attributable to the combined effects of greater foliage area and maximum crown conductance, lower stomatal sensitivity to high VPD, and higher transpiration demand relative to the control treatments. There was no significant interaction between CO2 and temperature on sap flow, because ECT entailed approximately similar patterns of sap flow to ET, suggesting that the temperature played a dominate role in the case of ECT under boreal climate conditions.

Carbon Dioxide↗

PubMed2005

Storage versus substrate limitation to bole respiratory potential in two coniferous tree species of contrasting sapwood width.

Two coniferous tree species of contrasting sapwood width (Pinus ponderosa L., ponderosa pine and Pseudotsuga menziesii Mirb., Douglas-fir) were compared to determine whether bole respiratory potential was correlated with available storage space in ray parenchyma cells and/or respiratory substrate concentration of tissues (total nitrogen content, N; and total non-structural carbohydrate content, TNC). An increment core-based, laboratory method under controlled temperature was used to measure tissue-level respiration (termed respiratory potential) from multiple positions in mature boles (>100-years-old). The most significant tissue-level differences that occurred were that N and TNC were two to six times higher for inner bark than sapwood, TNC was about two times higher in ponderosa pine than Douglas-fir and there was significant seasonal variation in TNC. Ray cell abundance was not correlated with sapwood respiratory potential, whereas N and TNC often were, implying that respiratory potential tended to be more limited by substrate than storage space. When scaled from cores to whole boles (excluding branches), potential net CO2 efflux correlated positively with live bole volume (inner bark plus sapwood), live bole ray volume, N mass, and TNC mass (adjusted R2 > or =0.4). This relationship did not differ between species for N mass, but did for live bole volume, live bole ray volume, and TNC mass. Therefore, N mass appeared to be a good predictor of bole respiratory potential. The differences in net CO2 efflux between the species were largely explained by the species' relative amounts of whole-bole storage space or substrate mass. For example, ponderosa pine's inner bark was thinner than Douglas-fir's, which had the greater concentration of ray cells and TNC compared with the sapwood. This resulted in ponderosa pine boles having 30-60% less ray volume and 10-30% less TNC mass, and caused ponderosa pine net CO2 efflux/ray volume and net CO2 efflux/TNC mass to be 20-50% higher than Douglas-fir. In addition, because inner bark respiratory potential was 2-25 times higher than that of sapwood, ponderosa pine's thinner inner bark and deeper sapwood (relative to Douglas-fir) caused its bole net CO2 efflux/live bole volume to be 20-25% lower than that of similarly-sized Douglas-fir trees.

Carbohydrate Metabolism↗

PubMed2006

Reducing stem bending increases the height growth of tall pines.

The hypothesis was tested that upper limits to height growth in trees are the result of the increasing bending moment of trees as they grow in height. The increasing bending moment of tall trees demands increased radial growth at the expense of height growth to maintain mechanical stability. In this study, the bending moment of large lodgepole pine (Pinus contorta Dougl. Ex Loud. var. latifolia Engelm.) was reduced by tethering trees at 10 m height to counter the wind load. Average bending moment of tethered trees was reduced to 38% of control trees. Six years of tethering resulted in a 40% increase in height growth relative to the period before tethering. By contrast, control trees showed decreased height growth in the period after tethering treatment. Average radial growth along the bole, relative to height growth, was reduced in tethered trees. This strongly suggests that mechanical constraints play a crucial role in limiting the height growth of tall trees. Analysis of bending moment and basal area increment at both 10 m and 1.3 m showed that the amount of wood added to the stem was closely related to the bending moment produced at these heights, in both control and tethered trees. The tethering treatment also resulted in an increase in the proportion of latewood at the tethering height, relative to 1.3 m height. For untethered control trees, the ratio of bending stresses at 10 m versus 1.3 m height was close to 1 in both 1998 and 2003, suggesting a uniform stress distribution along the outer surface of the bole.

Biomass↗