Lodgepole Pine Trees Research Articles

A specialized ABC efflux transporter GcABC‐G1 confers monoterpene resistance to Grosmannia clavigera, a bark beetle‐associated fungal pathogen of pine trees

Grosmannia clavigera is a bark beetle-vectored pine pathogen in the mountain pine beetle epidemic in western North America. Grosmannia clavigera colonizes pines despite the trees' massive oleoresin terpenoid defences. We are using a functional genomics approach to identify G. clavigera's mechanisms of adaptation to pine defences. We annotated the ABC transporters in the G. clavigera genome and generated RNA-seq transcriptomes from G. clavigera grown with a range of terpenes. We functionally characterized GcABC-G1, a pleiotropic drug resistance (PDR) transporter that was highly induced by terpenes, using qRT-PCR, gene knock-out and heterologous expression in yeast. Deleting GcABC-G1 increased G. clavigera's sensitivity to monoterpenes and delayed development of symptoms in inoculated young lodgepole pine trees. Heterologous expression of GcABC-G1 in yeast increased tolerance to monoterpenes. G. clavigera but not the deletion mutant, can use (+)-limonene as a carbon source. Phylogenetic analysis placed GcABC-G1 outside the ascomycete PDR transporter clades. G. clavigera appears to have evolved two mechanisms to survive and grow when exposed to monoterpenes: GcABC-G1 controls monoterpene levels within the fungal cells and G. clavigera uses monoterpenes as a carbon source. This work has implications for understanding adaptation to host defences in an important forest insect-fungal system, and potentially for metabolic engineering of terpenoid production in yeast.

Changes in transpiration and foliage growth in lodgepole pine trees following mountain pine beetle attack and mechanical girdling

The recent mountain pine beetle outbreak in North American lodgepole pine forests demonstrates the importance of insect related disturbances in changing forest structure and ecosystem processes. Phloem feeding by beetles disrupts transport of photosynthate from tree canopies and fungi introduced to the tree’s vascular system by the bark beetles inhibit water transport from roots to canopy; the implications of these processes for tree mortality are poorly understood. We hypothesized that the fungus must quickly disrupt tree water relations because phloem girdling, reported in other studies, requires much longer than a year to cause mortality. We tested the hypothesis in lodgepole pine (Pinus contorta) by comparing tree water use, foliar expansion and seasonal variation in predawn water potential on 8 mechanically girdled trees, 10 control trees and 17 trees attacked by mountain pine beetle (Dendroctonous ponderosae). Transpiration began to decline within ten days of beetle infestation; two months later, pre-dawn water potential had also dropped significantly as water transport to the canopy declined by 60% relative to healthy trees. There was no water transport or foliar expansion by beetle-infested trees the following year. Experimentally girdled trees continued to transpire, maintain leaf water potential and grow new foliage similar to healthy trees. Our data suggest that fungi introduced by bark beetles in this study are the primary cause of tree mortality following mountain pine beetle attack and significantly reduce transpiration soon after beetle infestation. Rapid decline and the eventual cessation of water uptake by infected trees have important implications for water and nutrient cycling in beetle impacted forests.

Variation in carbon availability, defense chemistry and susceptibility to fungal invasion along the stems of mature trees

If carbon (C) sinks withdraw carbohydrates as they are transported along tree stems, carbohydrate availability may depend on local sink strength and distance from sources. Defenses, including monoterpenes--a major component of resin--limit the invasibility of pines. Since carbohydrate reserves fund monoterpene synthesis, we hypothesized that monoterpene concentrations in pine stems would decrease from the crown to the lower stem, and susceptibility to fungal infection would increase. Here, we measured carbohydrate and monoterpene concentrations along the stems of lodgepole pine trees (Pinus contorta var. latifolia) before inoculating with a blue-stain fungus at different heights. After 6 wk, we assessed tree responses to fungal infection based on lesion length and carbohydrate mobilization. Concentrations of carbohydrates and monoterpenes in the phloem before inoculation decreased with distance from the crown, whereas lesion lengths after inoculation increased. However, trees mobilized sugars in response to fungal infection such that carbohydrate reserves near lesions were similar at all heights. Despite C mobilization, the lower stem was more vulnerable than the upper stem. Consistent with predictions based on sink-source relationships, vulnerability occurred where carbohydrates were less available, and likely resulted from C withdrawal by sinks higher in the supply chain.

Joint analysis of multivariate spatial count and zero‐heavy count outcomes using common spatial factor models

This paper discusses joint outcome modeling of multivariate spatial data, where outcomes include count as well as zero‐inflated count data. The framework utilized for the joint spatial count outcome analysis reflects that which is now commonly used for the joint analysis of longitudinal and survival data, termed shared frailty models, in which the outcomes are linked through a shared latent spatial random risk term. We discuss these types of joint mapping models and consider the benefits achieved through such joint modeling in the disease mapping context. We also consider the power of tests for common spatial structure in the context of two spatial maps and develop recommendations on the sort of power achievable in some contexts, as well as overall recommendations on the utility of joint mapping. We illustrate the approaches in an analysis of lung cancer mortality as well as an ecological study of Comandra blister rust infection of lodgepole pine trees. Copyright © 2012 John Wiley & Sons, Ltd.

The Impact of Phloem Nutrients on Overwintering Mountain Pine Beetles and Their Fungal Symbionts

In the low nutrient environment of conifer bark, subcortical beetles often carry symbiotic fungi that concentrate nutrients in host tissues. Although bark beetles are known to benefit from these symbioses, whether this is because they survive better in nutrient-rich phloem is unknown. After manipulating phloem nutrition by fertilizing lodgepole pine trees (Pinus contorta Douglas var. latifolia), we found bolts from fertilized trees to contain more living individuals, and especially more pupae and teneral adults than bolts from unfertilized trees at our southern site. At our northern site, we found that a larger proportion of mountain pine beetle (Dendroctonus ponderosae Hopkins) larvae built pupal chambers in bolts from fertilized trees than in bolts from unfertilized trees. The symbiotic fungi of the mountain pine beetle also responded to fertilization. Two mutualistic fungi of bark beetles, Grosmannia clavigera (Rob.-Jeffr. & R. W. Davidson) Zipfel, Z. W. de Beer, & M. J. Wingf. and Leptographium longiclavatum Lee, S., J. J. Kim, & C. Breuil, doubled the nitrogen concentrations near the point of infection in the phloem of fertilized trees. These fungi were less capable of concentrating nitrogen in unfertilized trees. Thus, the fungal symbionts of mountain pine beetle enhance phloem nutrition and likely mediate the beneficial effects of fertilization on the survival and development of mountain pine beetle larvae.

Effect of Bark Beetle Infestation on Secondary Organic Aerosol Precursor Emissions

Bark beetles are a potentially destructive force in forest ecosystems; however, it is not known how insect attacks affect the atmosphere. The emissions of volatile organic compounds (VOCs) were sampled i.) from bark beetle infested and healthy lodgepole pine (Pinus contorta var. latifolia) trees and ii.) from sites with and without active mountain pine beetle infestation. The emissions from the trunk and the canopy were collected via sorbent traps. After collection, the sorbent traps were extracted with hexane, and the extracts were separated and detected using gas chromatography/mass spectroscopy. Canister samples were also collected and analyzed by a multicolumn gas chromatographic system. The samples from bark beetle infested lodgepole pine trees suggest a 5- to 20-fold enhancement in total VOCs emissions. Furthermore, increases in the β-phellandrene emissions correlated with bark beetle infestation. A shift in the type and the quantity of VOC emissions can be used to identify bark beetle infestation but, more importantly, can lead to increases in secondary organic aerosol from these forests as potent SOA precursors are produced.

Association of tree diameter with body size and lipid content of mountain pine beetles

AbstractHost tree diameter is considered an important predictor of the population dynamics of the mountain pine beetle,Dendroctonus ponderosaeHopkins (Coleoptera: Curculionidae: Scolytinae), but the relationship between host tree diameter and beetle traits is unclear. The primary objective of this study was to determine how the quality of individual mountain pine beetles, measured as body size (pronotum width) and lipid content, varied with the diameter of lodgepole pine trees,Pinus contortavar.latifoliaEngelmann (Pinaceae). Naturally attacked trees, ranging in diameter from 10 to 35 cm, were selected from stands near Prince George, British Columbia, Canada. Colonisation density and pupal density generally increased with tree diameter, but the number of pupal chambers per gallery start remained constant. Tree diameter positively affected beetle body size, which in turn was positively correlated with absolute lipid content in both sexes and relative lipid content in males. However, tree diameter did not directly predict absolute lipid content, and relative lipid content decreased with tree diameter in males. Larger beetles emerged earlier in the emergence period with relative lipid content remaining constant throughout emergence. All relationships had considerable unexplained variation. Thus, the use of tree diameter or emergence time as predictors of population dynamics of mountain pine beetles should be done with caution.

Presence of carbaryl in the smoke of treated lodgepole and ponderosa pine bark

Lodgepole and ponderosa pine trees were treated with a 2% carbaryl solution at recreational areas near Fort Collins, CO, in June 2010 as a prophylactic bole spray against the mountain pine beetle. Bark samples from treated and untreated trees were collected one day following application and at 4-month intervals for one year. The residual amount of carbaryl was determined, and bark samples were burned to examine the smoke for the active ingredient. Smoke recovered from spiked bark samples showed a very high correlation between the treated rate and the concentration recovered from the smoke. Residual carbaryl on the bark was relatively stable throughout the study and carbaryl was detected in the smoke throughout the duration of the test.

Isolation and identification of diazotrophic bacteria from internal tissues ofPinus contortaandThuja plicata

Lodgepole pine ( Pinus contorta var. latifolia Engelm. ex S. Watson) and western red cedar ( Thuja plicata Donn ex D. Don) are capable of growing in soils that are severely nitrogen-limited. A rarely evaluated but possible nitrogen source for gymnosperms growing at nutrient poor sites is nitrogen fixation by endophytic bacteria. To investigate this possibility, we looked for nitrogen-fixing bacteria in surface-sterilized needle, stem, and root tissues of naturally regenerating lodgepole pine and western red cedar seedlings (2–4 years old) and trees (>20 years old) growing near Williams Lake, Chilliwack Lake, and Boston Bar, British Columbia, Canada. Ninety-nine bacterial isolates were obtained by plating surface-sterilized plant tissue extracts on tryptic soy agar (TSA) and combined carbon medium agar (CCMA), a nitrogen-free medium. Bacterial isolates were identified using gas chromatographic fatty acid methyl ester (GC-FAME) and 16S rRNA gene analyses and were evaluated for nitrogenase activity using an acetylene reduction assay. Representatives of the genera Bacillus and Paenibacillus dominated the culturable endophytic bacterial community in tissues of both tree species, with three Paenibacillus isolates and one Dyadobacter fermentans strain demonstrating consistently high acetylene-reduction activities. Our results raise the possibility that endophytic nitrogen fixation provides a supplementary nitrogen source for naturally regenerated lodgepole pine and western red cedar.

Variation in water potential, hydraulic characteristics and water source use in montane Douglas-fir and lodgepole pine trees in southwestern Alberta and consequences for seasonal changes in photosynthetic capacity

Tree species response to climate change-induced shifts in the hydrological cycle depends on many physiological traits, particularly variation in water relations characteristics. We evaluated differences in shoot water potential, vulnerability of branches to reductions in hydraulic conductivity, and water source use between Pinus contorta Dougl. ex Loud. var. latifolia Engelm. (lodgepole pine) and Pseudotsuga menziesii (Mirb.) Franco (interior Douglas-fir), and determined the consequences for seasonal changes in photosynthetic capacity. The Douglas-fir site had soil with greater depth, finer texture and higher organic matter content than soil at the lodgepole pine site, all factors that increased the storage of soil moisture. While the measured xylem vulnerability curves were quite similar for the two species, Douglas-fir had lower average midday shoot water potentials than did lodgepole pine. This implied that lodgepole pine exhibited stronger stomatal control of transpiration than Douglas-fir, which helped to reduce the magnitude of the water potential gradient required to access water from drying soil. Stable hydrogen isotope measurements indicated that Douglas-fir increased the use of groundwater during mid-summer when precipitation inputs were low, while lodgepole pine did not. There was a greater reduction of photosynthetic carbon gain in lodgepole pine compared with Douglas-fir when the two tree species were exposed to seasonal declines in soil water content. The contrasting patterns of seasonal variation in photosynthetic capacity observed for the two species were a combined result of differences in soil characteristics at the separate sites and the inherent physiological differences between the species.

Effect of Water Stress and Fungal Inoculation on Monoterpene Emission from an Historical and a New Pine Host of the Mountain Pine Beetle

The mountain pine beetle (Dendroctonus ponderosae, MPB) has killed millions of lodgepole pine (Pinus contorta) trees in Western Canada, and recent range expansion has resulted in attack of jack pine (Pinus banksiana) in Alberta. Establishment of MPB in the Boreal forest will require use of jack pine under a suite of environmental conditions different from those it typically encounters in its native range. Lodgepole and jack pine seedlings were grown under controlled environment conditions and subjected to either water deficit or well watered conditions and inoculated with Grosmannia clavigera, a MPB fungal associate. Soil water content, photosynthesis, stomatal conductance, and emission of volatile organic compounds (VOCs) were monitored over the duration of the six-week study. Monoterpene content of bark and needle tissue was measured at the end of the experiment. β-Phellandrene, the major monoterpene in lodgepole pine, was almost completely lacking in the volatile emission profile of jack pine. The major compound in jack pine was α-pinene. The emission of both compounds was positively correlated with stomatal conductance. 3-Carene was emitted at a high concentration from jack pine seedlings, which is in contrast to monoterpene profiles of jack pine from more southern and eastern parts of its range. Fungal inoculation caused a significant increase in total monoterpene emission in water deficit lodgepole pine seedlings right after its application. By 4weeks into the experiment, water deficit seedlings of both species released significantly lower levels of total monoterpenes than well watered seedlings. Needle tissue contained lower total monoterpene content than bark. Generally, monoterpene tissue content increased over time independent from any treatment. The results suggest that monoterpenes that play a role in pine-MPB interactions differ between lodgepole and jack pine, and also that they are affected by water availability.

Climate impacts on lodgepole pine ( Pinus contorta) radial growth in a provenance experiment

A potentially confounding radial-growth interaction exists at the intersection of two well-known principles, one in the field of dendrochronology and the other in quantitative genetics. From a dendrochronology perspective, tree populations growing in climatically marginal environments are expected to be more sensitive to seasonal and annual climate than those growing in optimal climate zones. From a genetics perspective, marginal populations may be adapted to grow a small amount each year and then shut down to prevent climate-induced mortality, or they may be adapted to respond to favourable climate conditions when available. We examined the relative strength of these forces using data from 12 populations of 34-year-old lodgepole pine ( Pinus contorta) trees growing in 16 provenance-trial sites in western Canada. Growth generally correlated positively with annual temperature and negatively with summer aridity. The sensitivity of radial-growth to interannual climate fluctuations was both site and provenance-related, with the highest sensitivities occurring among populations from warm, central provenances growing at cold, marginal sites, and among populations from cold, marginal provenances growing at warm, central sites. The correlations between climate and growth varied regionally; notably, populations from warm provenances growing at warm sites responded more strongly to summer aridity, while populations from cold provenances growing at warm sites responded more to annual temperature. Our finding that sensitivity varied among populations growing under similar climate conditions indicates that sensitivity is influenced by genetics as well as by site climate, but the regional specificity of the growth responses did not support a single hypothesis for the influence of genetics on growth among populations from marginal vs. central locations. Implications of our study for forest productivity under climate change are more positive for trees growing in cool locations, where overall warmer temperatures will lead to increased growth, than in warm locations, where the negative effects of arid summers may counteract the positive effects of warmer annual temperatures.

Modeling lodgepole pine radial growth relative to climate and genetics using universal growth-trend response functions

Forests strongly affect Earth's carbon cycles, making our ability to forecast forest-productivity changes associated with rising temperatures and changes in precipitation increasingly critical. In this study, we model the influence of climate on annual radial growth using lodgepole pine (Pinus contorta) trees grown for 34 years in a large provenance experiment in western Canada. We use a random-coefficient modeling approach to build universal growth-trend response functions that simultaneously incorporate the impacts of different provenance and site climates on radial growth trends under present and future annual (growth-year), summer, and winter climate regimes. This approach provides new depth to traditional quantitative genetics population response functions by illustrating potential changes in population dominance over time, as well as indicating the age and size at which annual growth begins declining for any population growing in any location under any present or future climate scenario within reason, given the ages and climatic conditions sampled. Our models indicate that lodgepole pine radial-growth levels maximize between 3.9 degrees and 5.1 degrees C mean growth-year temperature. This translates to productivity declining by the mid-21st century in southern and central British Columbia (BC), while increasing beyond the 2080s in northern BC and Yukon, as temperatures rise. Relative to summer climate indices, productivity is predicted to decline continuously through the 2080s in all locations, while relative to winter climate variables, the opposite trend occurs, with the growth increases caused by warmer winters potentially offsetting the summer losses. Trees from warmer provenances, i.e., from the center of the species range, perform best in nearly all of our present and future climate-scenario models. We recommend that similar models be used to analyze population growth trends relative to annual and intra-annual climate in other large-scale provenance trials worldwide. An open-access growth-trend data set encompassing numerous biomes, species, and provenances would contribute substantially to predicting forest productivity under future-climate scenarios.

Nitrogen cycling following mountain pine beetle disturbance in lodgepole pine forests of Greater Yellowstone

Widespread bark beetle outbreaks are currently affecting multiple conifer forest types throughout western North America, yet many ecosystem-level consequences of this disturbance are poorly understood. We quantified the effect of mountain pine beetle ( Dendroctonus ponderosae) outbreak on nitrogen (N) cycling through litter, soil, and vegetation in lodgepole pine ( Pinus contorta var. latifolia) forests of the Greater Yellowstone Ecosystem (WY, USA) across a 0–30 year chronosequence of time-since-beetle disturbance. Recent (1–4 years) bark beetle disturbance increased total litter depth and N concentration in needle litter relative to undisturbed stands, and soils in recently disturbed stands were cooler with greater rates of net N mineralization and nitrification than undisturbed sites. Thirty years after beetle outbreak, needle litter N concentration remained elevated; however total litter N concentration, total litter mass, and soil N pools and fluxes were not different from undisturbed stands. Canopy N pool size declined 58% in recent outbreaks, and remained 48% lower than undisturbed in 30-year old outbreaks. Foliar N concentrations in unattacked lodgepole pine trees and an understory sedge were positively correlated with net N mineralization in soils across the chronosequence. Bark beetle disturbance altered N cycling through the litter, soil, and vegetation of lodgepole pine forests, but changes in soil N cycling were less severe than those observed following stand replacing fire. Several lines of evidence suggest the potential for N leaching is low following bark beetle disturbance in lodgepole pine.

Diversity and decay ability of basidiomycetes isolated from lodgepole pines killed by the mountain pine beetle

When lodgepole pines (Pinus contorta Douglas ex Louden var. latifolia Engelm. ex S. Watson) that are killed by the mountain pine beetle (Dendroctonus ponderosae) and its fungal associates are not harvested, fungal decay can affect wood and fibre properties. Ophiostomatoids stain sapwood but do not affect the structural properties of wood. In contrast, white or brown decay basidiomycetes degrade wood. We isolated both staining and decay fungi from 300 lodgepole pine trees killed by mountain pine beetle at green, red, and grey stages at 10 sites across British Columbia. We retained 224 basidiomycete isolates that we classified into 34 species using morphological and physiological characteristics and rDNA large subunit sequences. The number of basidiomycete species varied from 4 to 14 species per site. We assessed the ability of these fungi to degrade both pine sapwood and heartwood using the soil jar decay test. The highest wood mass losses for both sapwood and heartwood were measured for the brown rot species Fomitopsis pinicola and the white rot Metulodontia and Ganoderma species. The sap rot species Trichaptum abietinum was more damaging for sapwood than for heartwood. A number of species caused more than 50% wood mass losses after 12 weeks at room temperature, suggesting that beetle-killed trees can rapidly lose market value due to degradation of wood structural components.

Phosphorus release from forest harvesting on an upland blanket peat catchment

The aim of this study was to investigate the release of phosphorus (P) to receiving waters resulting from harvesting 34-year-old lodgepole pine trees in an upland peat catchment. The study site was within a 25.3-hectare (ha) area, and was drained by a stream that received flows from ploughed furrows, mainly, via collector drains, and discharged directly to the salmonid Shrahrevagh River, Burrishoole, Co. Mayo, Ireland. The study site was divided into two parts: the upstream part was left intact and the downstream part was harvested in early Autumn 2005 following implementation of forest guidelines. Good management practices such as proper use of brash mats and harvesting only in dry weather were implemented. Two instrumented stations were established – one just upstream (US) and the other just downstream (DS) of the clearfelled area. The measurement of P concentrations at the two stations commenced in May 2005, two months before the harvesting started. The daily mean P concentration at the DS station increased from about 6 μg L −1 of total reactive phosphorus (TRP) during pre-clearfelling to 429 μg L −1 in August 2006. By October 2009, four years after clearfelling, the P concentrations at the DS station had returned to pre-clearfelling levels. In the first three years after harvesting, up to 5.15 kg ha −1 of TRP was released from the harvested catchment to the receiving water; in the second year alone, 2.3 kg ha −1 of TRP was released. Linear regression can be used to describe the relationship between TRP load and water discharge. About 80% of the total phosphorus (TP) in the study stream was soluble and more than 70% of the P release occurred in storm events, indicating that traditional buffer strips with widths of 15–20 m might not be efficient for P immobilization. The P concentrations were affected by antecedent weather conditions and highest concentrations occurred during storm events following prolonged drought periods. The water extractable phosphorus (WEP) contents in the soil were significantly higher below windrow/brash material than in brash-free areas, and whole-tree harvesting should be studied as one of the means to decrease P export from blanket peats.

ForestSimMPB: A swarming intelligence and agent-based modeling approach for mountain pine beetle outbreaks

The widespread outbreaks of Mountain Pine Beetle (MPB) are responsible for infestations of lodgepole pine forests since 1990 in Canada. In British Columbia, this forest insect disturbance has resulted in losses of more than 13 million hectares of pine trees. The complexity of the MPB emergence, aggregation and attack behaviour is captured by this study, using an intelligent agent-based model (ABM) of beetle outbreaks at a local scale of individual trees. Agent-based approach permits simulation of interactions that describe the ecological context in which insect populations spread. Intelligent reasoning is introduced by a swarm intelligence (SI) algorithm integrated with the ABM that depicts indirect communication, collective behaviour and self-organized aggregation of insects in a forest ecosystem. The objectives of this study are the following: 1) to develop ForestSimMPB model that integrates SI and ABM within a geographic information systems (GIS) framework; 2) to implement the proposed model on real datasets to simulate the MPB aggregation and mass attacks on lodgepole pine trees; and 3) to determine the spatial patterns and extents of these attacks. The ForestSimMPB is calibrated by fine tuning two model parameters, and implemented using data from three sites located in the Cariboo Regional District in the central interior of BC. The obtained results demonstrate the aggregation behaviour of MPB to collectively attack lodgepole pines, as well as portray the spatial clustering of dead trees resulting from infestation. Simulation outputs provide analysis and predictions of spatial patterns in the forest landscape structure as a result of a MPB outbreak. The developed model can be used to assist the improvement of methods for prevention and control of MPB disturbances.

A Gamma-Poisson model for vertical location and frequency of buds on lodgepole pine (Pinus contorta) leaders

The aim of this work was to model the vertical location and number of branch primordia (buds) on the leader of lodgepole pine ( Pinus contorta Doug. ex Loud.) trees in central British Columbia. For species such as lodgepole pine, where branches occur in clusters rather than individually, the Gamma-Poisson model provides a natural framework for describing and simulating the distribution of buds on the annual shoot. Parameters in the model are identifiable with measurable attributes, that is, the average number of clusters per unit length of the annual shoot and the average number of buds per cluster, and can be related to explanatory variables via a log link. Applicability of the Gamma-Poisson model was demonstrated for a sample of 58 lodgepole pine trees ranging in age from 29 to 103 years old. The agreement between observed and expected cluster counts and spacing, cluster sizes, and total number of branches was good. Height to crown base and length of the annual shoot were selected as the best predictors of the number of clusters and number of buds per cluster, respectively, although other single variables were also identified as having significant predictive value.

Fertilization of lodgepole pine trees increased diameter growth but reduced root carbohydrate concentrations

Tree growth variables and more complex derived variables such as vigor index have all been linked to the ability of lodgepole pine (Pinus contorta var latifolia) trees to defend against insect herbivores, particularly mountain pine beetle (Dendroctonus ponderosae Hopkins, Coleoptera: Curculionidae). These variables are considered indirect measures of carbohydrate reserves. Trees with high vigor index values have high growth rates and are assumed to have high carbohydrate reserves. However, this critical assumption was untested for lodgepole pine. In this paper we evaluated the relationship between carbohydrate concentration and tree characteristics. We measured concentrations of root starch in 10 pure lodgepole pine stands of different ages in the Canadian Rockies which had been fertilized, thinned, fertilized and thinned or left untreated in a 2×2 factorial design. Both fertilization and thinning increased basal area increment while only fertilization increased lateral branch growth in the crown. Contrary to expectations, fertilization decreased root starch reserves although it increased basal area increment; thinning had no effect on root carbohydrates. Root starch reserves were positively related to basal area increment and vigor index in both fertilized and unfertilized plots, however, the best predictors of starch reserves were height-to-live-crown and cambial surface area below the live crown; starch reserves declined as both of these variables increased. These data suggest that large carbon sinks associated with long boles below the crown, and rapid growth of the crown and stem as a result of fertilization are detrimental to building starch reserves in the roots.

Evaluation of Mountain Beetle-Infested Lodgepole Pine for Cellulosic Ethanol Production by Sulfite Pretreatment to Overcome Recalcitrance of Lignocellulose

The potentials of deteriorated mountain pine beetle (Dendroctonus ponderosae)-killed lodgepole pine (Pinus contorta) trees for cellulosic ethanol production were evaluated using the sulfite pretreatment to overcome recalcitrance of lignocellulose (SPORL) process. The trees were harvested from two sites in the United States Arapaho-Roosevelt National Forest, Colorado. The infestation age of the trees varied from zero to about 8 years. Mild (170 °C) and harsh (180 °C) SPORL pretreatments were conducted. The chemical charges were sulfuric acid of 2.21% and sodium bisulfite of 8% on oven dry wood for the harsh and half of those for the mild pretreatment. The results suggest that beetle-caused mortality enriched glucan content by as much as 3 percentage points (or 7.5%) in wood. The glucan enrichment seems to increase with infestation age. The enriched glucan can be captured after SPORL pretreatment followed by enzymatic hydrolysis. The killed trees are more susceptible to SPORL pretreatment, which enhanced substrate enzymatic digestibility (SED). Enzymatic hydrolysis glucose yields (EHGY) from killed trees were about 5-20% higher than those from their corresponding live trees. Total fermentable sugar productions from dead trees (including a tree laying on the ground) were 4-14% higher than corresponding production from live trees, depending on pretreatment conditions and infestation age. An ethanol yield of 267 L/metric ton of wood or 69% theoretical value was achieved from a tree infested 4 years, 7% higher than the 250 L/metric ton of wood from the corresponding live tree. The results also demonstrated the robustness of SPORL pretreatment for lodgepole pine.