Pinus Monticola Dougl Research Articles

Soil disturbance, amelioration and rehabilitation affect forest growth, health, soil carbon and chemistry on five long-term soil productivity (LTSP) sites in southeastern British Columbia

Soil disturbance amelioration affected tree survival, growth, health, mineral soil chemistry, and their covariation on five Long-Term Soil Productivity (LTSP) sites in southeastern British Columbia. The two most different LTSP treatments (baseline disturbance of no compaction / all organic matter left, versus most severe of heavy compaction / forest floor removed) were compared to five amelioration treatments: rehabilitation of most severe LTSP treatment (soil tillage with or without incorporation of removed forest floor [Rehab + FF]), root disease (Armillaria ostoyae (Romagn.) Herink) amelioration treatments (stump removal or biological with Hypholoma fasiculare), and enhancement of soil organic matter (tilling extra forest floor into mineral soil). Treatment significantly (p < 0.05) affected 10-year mineral soil carbon, nitrogen, sulfur, mineralizable nitrogen, and potassium, which were lower on forest floor removal treatments and higher on forest floor addition treatments (27 to 97% of baseline versus 78 to 183% of baseline, respectively). Western white pine (Pinus monticola Dougl. ex D.Don) 15-year productivity (m3/ha) was significantly affected by treatment (Rehab + FF yielding a 10 fold increase in productivity versus the poorly growing baseline treatment), with treatment also significant when total disease was a covariate and for average and total volume when mineral soil Ca:Al ratio was a marginally significant (p < 0.1) covariate, and for survival when the incidence of total disease or white pine blister rust (Cronartium ribicola) were significant covariates; or with mineral soil C:N ratio as a significant covariate. The incidence of blister rust and total disease were significantly influenced by treatment when C:N was a significant covariate. For 20-year lodgepole pine (Pinus contorta Dougl. Ex Loud) there were no direct relationships apparent, but western gall rust (Endocronartium harknessii) had treatment significant when total sulfur was a significant covariate. Fifteen- and twenty-year Douglas-fir (Pseudotsuga menziesii var. glauca [Beissn.] Franco) had no significant relationships. Results demonstrate that restoring or adding extra organic matter (forest floor) into mineral soil increased soil carbon and related nutrients and can increase above ground carbon (forest productivity). Conversely, removing forest floor lowered mineral soil carbon and generally lowered tree volume. Varied effects due to growth limiting factors like frost and competing vegetation affected results on a few plots. Recommendations include leaving or enhancing organic matter onsite for soil carbon, nutrients, forest health and site productivity. Further analysis with the larger LTSP dataset and ongoing monitoring is encouraged to determine if the results persist in the long-term and to better understand the significant covariate relationships.

White pine blister rust, logging, and species replacement increased streamflow in a montane watershed in the northern Rockies, USA

Vegetation changes can strongly influence the hydrological cycle, including streamflow, but the effects of plant diseases have seldom been described. In the mid-20th century, the invasion of an exotic disease, white pine blister rust, precipitated widespread mortality of western white pine (WWP; Pinus monticola Dougl.) in the northern Rocky Mountains, USA. These events converted a forest dominated by white pine into one dominated by more shade-tolerant tree species. The long-term hydrological implications of this historical shift in forest composition have not been adequately explored, in part because collocated long-term vegetation, meteorology, and hydrology data are rare. We assembled long-term streamflow, climate, and vegetation records for a small (4.0 km2) forested watershed in the U.S. northern Rocky Mountains. We used a minimally-calibrated, physically-based model to simulate historical changes in the water budget based on observed vegetation changes along with the historical climate. The observed hydrological anomalies were attributed to a combination of white pine blister rust induced tree mortality, harvest, and eventual species replacement. Small increases in streamflow began in the basin in the 1940s, when the basal area of WWP began to decrease. More dramatic increases in streamflow started after harvesting began in 1966. Based on both observations and modeling, streamflow increased by ∼131–179 mm between the 1940s and the 2000s (30–40% of the 2000s’ streamflow) in contrast to documented regional streamflow declines. Approximately 1/3 of the flow changes were attributed to blister rust and 2/3 to harvest, as shade-tolerant species instead of WWP regenerated in response to both. This study highlights the importance of long-term ecohydrological data, which made it possible to detect the influence of the invasion of an exotic pathogen against a backdrop of periodic timber harvest. It also showed that the species replacement after these disturbances caused long-term changes in the flow regime, providing a clear and relevant example of how land cover changes can affect interannual hydrological dynamics.

Fifteen-year tree growth on standard long-term soil productivity trials and various adjacent amelioration treatments at Interior Cedar-Hemlock sites in southeastern British Columbia and northern Idaho

Understanding the relationship between soil disturbance and tree growth is key to sustainable forest management. We report fifteen-year tree growth results of Douglas-fir (Pseudotsuga menziesii var. glauca [Beissn.] Franco) and western white pine (Pinus monticola Dougl. ex D.Don) growth following clearcut harvesting and installation of three replicates of the international Long-Term Soil Productivity (LTSP) trial protocols. Additional amelioration plots were also established on each replicate site to study rehabilitation of the most severe disturbance, treatment of Armillaria root disease, and enhancement of soil organic matter. Stark differences were found among the various treatments and species. Vegetation control resulted in significantly better growth for both tree species across the core LTSP treatments and amelioration treatments. On the core LTSP treatments, organic matter removal had a significant negative effect on Douglas-fir growth. Amelioration treatments resulted in similar survival and similar or better volume growth than the core LTSP treatments, with the exception of poor volume growth after compaction rehabilitation with no organic matter returned into the soil. Compaction significantly influenced Douglas-fir survival and was often higher on moderate compaction plots. On some sites there was decreased survival and growth associated with heavy compaction that will likely influence longer-term outcomes with a more variable and changing climate. Overall, results to date on the core LTSP treatments, amelioration treatments, and adjacent miniplots reported elsewhere suggest that forest floor removal causes negative growth and, combined with severe compaction, caused both negative growth and poor survival. In the Interior Northwest it is still prudent to minimize soil disturbance and leave organic matter onsite unless there is site specific evidence to the contrary. Additional work is required to reap the full, long-term benefit of the installations. Recommendations are provided highlighting the critical work that should be completed in the near future before biological evidence is lost to decay, including suggestions for further data analysis and extension, the need to continue LTSP core, amelioration and miniplot measurements to demonstrate and calibrate sustainable forest resource management. Our data suggest considering changes to current soil disturbance policy and forest operations based on the current results regarding forest floor displacement.

Evaluation of Stem-Injected TREE-äge® (4% Emamectin Benzoate) for Protecting Western White Pines (Pinus monticola) from Mountain Pine Beetle (Dendroctonus ponderosae Hopkins)(Coleoptera: Curculionidae: Scolytinae)

The protection of high-value trees against bark beetles and the development of alternatives to bole sprays is a priority for the tree manager. The objective of this study was to evaluate stem-injected TREE-äge® (emamectin benzoate [EB]) as a protective treatment for western white pines (Pinus monticola Dougl. ex D. Don) against mountain pine beetle (MPB, Dendroctonus ponderosae Hopkins). Treatment efficacy was based solely on tree mortality as per Shea protocols (i.e., ≥ 60% check vs. ≤ 20% treated tree mortality). Our first experiment was installed in 2007 and included trees stem-injected with TREE-äge and untreated controls. Bole application of S-(-)-verbenone and green leaf volatile (GLV) blend was included for observational comparison. Pressure from MPB was heavy, as indicated by the number and timing of control tree mortality (90%). Strip attacks by MPB in TREE-äge trees indicated that the impacts of EB, and by inference its distribution, were inconsistent. In 2009, the injection protocol was revised to improve EB distribution in the phloem via closer injection points. In the 2009 TREE-äge-treated trees, adult beetle mining stopped when they contacted phloem and was insufficient to cause tree death by girdling. Blue-stain fungi colonized the sapwood of trees in both studies. Isolates from autopsied trees treated with TREE-äge alone were subsequently identified as Grosmannia clavigera and Leptographium longiclavatum (Ophiostomatales: Ascomycota), species that can incite tree mortality. In 2013, we revised our protocol to include GLV plus verbenone or propiconazole with TREE-äge, wherein these treatments proved effective in protecting trees against MPB and their associated pathogenic fungi.

Transient gene expression in western white pine using agroinfiltration

A method for transient gene expression was developed for western white pine (WWP, Pinus monticola Dougl. ex D.Don) using reporter gene uidA encoding β-glucuronidase (GUS). GUS was transiently expressed in cross sections of primary and secondary needles, cotyledons, and current and second year stems of WWP via vacuum-infiltration with Agrobacterium tumefaciens. Histochemical assays of cross sections of secondary needles showed stronger blue color indicating GUS expression at day 1 and 2 than on other days post agroinfiltration (dpa). GUS activity expressed inside WWP cells was confirmed using light microscopy. In fluorometric assays, GUS expression was high at 1 dpa and lasted until 4 dpa in detached secondary needles, while similarly high expression levels only lasted until 2 dpa in attached secondary needles then dropped significantly. Although the length of GUS-staining zones varied among different WWP organs and between growth and dormant seasons, all tested WWP tissues using the protocol had high levels of transient GUS expression. Thus, heterologous candidate genes or endogenous silencing can be expressed in various WWP tissues or organs using this agroinfiltration approach. The current protocol for efficient transient gene expression will aid functional genomics study of WWP and its pathogens and related conifer species.

Adaptive response of Pinus monticola driven by positive selection upon resistance gene analogs (RGAs) of the TIR-NBS-LRR subfamily

Abstract: Western white pine (Pinus monticola Dougl.) is an important forest tree species, which is intensively plagued by the fungus Cronartium ribicola. Resistance gene analogs (RGAs) are the most highly abundant class of potential resistance (R) genes sharing greatly conserved domains and structures. Hence RGAs are crucial components for disease resistance breeding programs on P. monticola serving as useful functional markers. A total of 33 P. monticola RGAs gene homologues were mined from GenBank, encoding for R gene members of the TIR-NBS-LRR subfamily. The existence of positive selection acting upon RGAs was determined using a series of maximum likelihood analyses. Robust evidence of positive selection was showed to be acting widely in three clades across RGA gene phylogeny, both on terminal and ancestral lineages. Furthermore, our analysis revealed that the majority of positively selected residues sites are localized widely across these RGAs sequences, putatively affecting the structures of their ligand-binding domains and offering novel specificities. These results may find immediate application in ongoing disease resistance breeding programs.

Sulfometuron methyl influences seedling growth and leaf function of three conifer species

Seedling growth and gas exchange responses were measured for two potted seedling trials testing herbicide phytotoxicity to three important tree species of the Inland Northwest, USA. Media-filled pots were treated with sulfometuron methyl (Oust®) in varying concentrations and planted with seedlings of Larix occidentalis Nutt., Pseudotsuga menziesii var. glauca (Beissn.) Franco, and Pinus monticola Dougl. ex D. Don. Seedlings were grown in two trials to determine the effects of two important residue breakdown factors, substrate moisture and pH, relative to that of herbicide application rate on seedling health. Changes in seedling height, root-collar diameter, and root volume were morphologic variables of interest, and physiological variables measured were net photosynthetic assimilation, stomatal conductance, and transpiration rate. While three levels of media moisture and four levels of media pH had no effect on seedling performance, most growth and leaf function variables were hindered across application rate treatments of all three species. Label-recommended dosages resulted in growth suppression levels potentially detrimental to seedling establishment for all three species. This was most pronounced for Pinus monticola, in particular for root growth, where untreated control seedlings showed 109% more root volume growth than treated seedlings. We conclude that when possible, a species-specific application rate might be found that balances the benefits of vegetation control with the phytotoxicity to promote optimal growth gains.

Tree growth at stand and individual scales in two dual-species mixture experiments in southern Washington State, USA

Planting with mixtures of tree species rather than single species is often considered during reforestation because of the potential increased productivity and other benefits. We examined tree growth at the stand and individual tree scales in two experiments contrasting monocultures with a 1:1 mixture of tree species: (1) Douglas-fir ( Pseudotsuga menziesii (Mirb.) Franco) with a conifer of similar shade tolerance (western white pine ( Pinus monticola Dougl. ex D. Don)) and (2) Douglas-fir with a more shade-tolerant conifer (western hemlock ( Tsuga heterophylla (Raf.) Sarg.)). There was no effect of mixture on growth or yield in the Douglas-fir – western white pine combination. In the Douglas-fir – western hemlock combination, yields in the mixture equaled those in Douglas-fir stands because of the enhanced performance of Douglas-fir in the mixture. For Douglas-fir, the height/diameter (h/d) ratio was significantly less in mixture, suggesting reduced competition for light when grown with western hemlock. In contrast, the h/d ratio for western hemlock was significantly greater in mixture, suggesting increased competition for light when grown with Douglas-fir. Neighborhood analyses showed that tree growth was directly related to initial size and inversely related to relative neighbor size and that the h/d ratio was positively related to relative neighbor size. In general, the size of a neighboring tree influenced growth more than species identity. Relationships between h/d ratios and growth rates suggest that growth differences between Douglas-fir and western hemlock in mixture will eventually increase.

Crown Recession Patterns in Three Conifer Species of the Northern Rocky Mountains

Abstract Crown length is a fundamental tree dimension for characterizing growth potential, wildlife habitat, and wood quality. The relative rates of height growth and crown recession determine the progression of crown length over time. We investigated patterns in crown recession of three co-occurring species in the northern Rocky Mountains: western white pine (Pinus monticola Dougl. ex D. Don), ponderosa pine (Pinus ponderosa Dougl. ex P. &amp; C. Laws.), and interior Douglas-fir (Pseudotsuga menziesii [Mirb.] Franco var. glauca). Past height, diameter, and crown base on 169 forest-grown trees were reconstructed by detailed stem analysis. Crown base was estimated for each year of the tree's life by dating the mortality of all whorl branches. Five-year crown recession was modeled in two parts, the first predicting the probability that a crown recedes and the second estimating recession conditional on its occurrence. The probability of crown recession increased and then decreased with an increasing crown ratio for all three species and similarly peaked over initial tree size. Conditional crown recession increased monotonically with crown ratio for ponderosa pine but peaked at crown ratios between 0.7 and 0.9 in western white pine and Douglas-fir. The resulting models lend insight into factors controlling rates of crown recession and their variation among species and differing initial conditions.

The coat-enhanced dormancy mechanism of western white pine (Pinus monticola Dougl. ex D. Don) seeds is mediated by abscisic acid homeostasis and mechanical restraint

The coat-enhanced dormancy mechanism of western white pine (Pinus monticola Dougl. ex D. Don) seeds is mediated by abscisic acid homeostasis and mechanical restraint

First Report of the White Pine Blister Rust Fungus, Cronartium ribicola, on Pedicularis bracteosa.

Until recently, Cronartium ribicola J.C. Fisch. was thought to utilize only Ribes spp. (Grossulariaceae) as telial hosts in North America. During 2004, Pedicularis racemosa Dougl. ex Benth. and Castilleja miniata Dougl. (Orobanchaceae) were proven as natural telial hosts at a subalpine site (48.634109°N, 116.570817°W, elevation 1,800 m) near Roman Nose Lake, ID, where whitebark pine (Pinus albicaulis Engelm.) and western white pine (Pinus monticola Dougl. ex D. Don) are aecial hosts, and Pedicularis, Castilleja, and Ribes spp. are common herbs/shrubs (2). During August 2006, teliospore columns typical of C. ribicola or the morphologically indistinguishable (2) C. coleosporioides J.C. Arthur were found on two Pedicularis bracteosa Benth. plants at this site, within 3 m of a large, sporulating canker on whitebark pine. ITS/5.8S rDNA regions were sequenced using detached teliospore column samples from the two plants, ITS1F and ITS4 primers (3), and standard PCR protocols (2). One sample sequence was identified as C. ribicola and the other as C. coleosporioides (GenBank Accession Nos. EF185857 and EF185858, respectively), by exact matches in comparisons with published sequences (2). Artificial inoculation confirmed P. bracteosa's ability to host C. ribicola. Sections of leaves collected near Freezeout Saddle, ID (47.00885°N, 116.00846°W, elev. 1,600 m) were rinsed in water, placed abaxial side up on moistened filter paper in 150-mm petri plates, inoculated with seven diverse sources of urediniospores/aeciospores, misted with distilled water, and incubated at 18°C with 12 h of light. A single leaf section produced urediniospores 17 days and teliospores 26 days after inoculation with one of two Roman Nose aeciospore sources. Urediniospores from this leaf section caused infections on Ribes nigrum L., and teliospore columns yielded a DNA sequence that matched C. ribicola. Though P. bracteosa is confirmed as yet another natural host of C. ribicola in North America, it may be producing less C. ribicola inoculum for pine infection than do the P. racemosa and Ribes spp. telial hosts at the collection site. Uredinia and telia of C. ribicola on P. bracteosa were much less frequent and smaller than those on P. racemosa and Ribes spp. and those of C. coleosporioides on this same host (2). Pedicularis (but not Castilleja) spp. are significant telial hosts of C. ribicola strains at some high elevation sites in eastern Asia (1). Discovery of multiple North American telial hosts in the Orobanchaceae suggests unrecognized complexity in C. ribicola's ability to exploit ecological niches in recently established pathosystems of North America (2).

Pollination and seed production in western white pine

The reproductive biology of western white pine ( Pinus monticola Dougl. ex D. Don) at the Saanich seed orchard (SSO; a coastal seedling seed orchard) and the Kalamalka seed orchard (KSO; an interior clonal seed orchard) is described. Seed-cone development and morphology determine seed potential and affect filled seed production. Seed potential was high, commonly over 200 seeds per cone and filled seeds per cone averaged 50–60 at both orchards in 1999 from open pollinations combined with operational supplemental mass pollinations (OP-SMP). Filled seeds per cone at SSO in 2004 averaged 115 with open pollinations and OP-SMP. About 30% of ovules aborted before pollination. Another 25% aborted probably because of self-incompatibility at fertilization. Seed efficiency (SEF) was only 5% at KSO with open pollination but increased to 30% with OP-SMP. SEF was the same at SSO with or without OP-SMP. Reproductive success was very low at KSO without OP-SMP but increased to about 10%, similar to that of SSO, with OP-SMP. The optimal time for control pollinations was at cone stages 4 and 5 and optimal amount of pollen per pollination bag was 0.3–0.4 mL. Western white pine is a very good seed producer and can be easily managed in seed orchards to give very high seed production per cone and per tree.

Co-occurring species differ in tree-ring 18O trends

The stable oxygen isotope ratio (delta(18)O) of tree-ring cellulose is jointly determined by the delta(18)O of xylem water, the delta(18)O of atmospheric water vapor, the humidity of the atmosphere and perhaps by species-specific differences in leaf structure and function. Atmospheric humidity and the delta(18)O of water vapor vary seasonally and annually, but if the canopy atmosphere is well mixed, atmospheric characteristics should be uniform among co-occurring trees. In contrast, xylem water delta(18)O is determined by the delta(18)O of water being drawn from the soil, which varies with depth. If co-occurring trees draw water from different soil depths, this soil-water delta(18)O signal would be manifest as differences in delta(18)O among the trees. We examined the variation in tree ring delta(18)O, over eight decades during the 20th Century, among three species co-occurring in natural forest stands of the northern Rocky Mountains in the USA. We sampled 10 Douglas-firs (Pseudotsuga menziesii (Mirb.) Franco var. glauca), 10 ponderosa pines (Pinus ponderosa Laws.) and seven western white pines (Pinus monticola Dougl.). As expected, variation in atmospheric conditions was recorded in the delta(18)O of the cellulose produced in a given year, but observed climatic correlations with delta(18)O were weak. Significant correlations with June climate data included: daily maximum temperature (r = 0.29), daily minimum temperature (r = -0.25), mean temperature (r = 0.20), mean daily precipitation (r = -0.54), vapor pressure deficit (r = 0.32) and solar radiation (r = 0.44). Lagged effects were observed in Douglas-fir and western white pine. In these species, the delta(18)O of a given annual ring was correlated with the delta(18)O of the previous ring. Ponderosa pine showed no significant autocorrelation. Although the species means were correlated among years (r = 0.67 to 0.76), ponderosa pine was consistently enriched in delta(18)O relative to the other species; differences were close to 2 per thousand and they are steadily increasing. Relative to the mean for the three species, ponderosa pine is becoming steadily more enriched (-1.0 per thousand). In contrast, Douglas-fir is being steadily depleted and western pine is intermediate, with an enrichment of 0.5 per thousand. Because all trees were exposed to the same atmospheric conditions, the differences in delta(18)O observed between species are likely due either to differences in the depth of water extraction or leaf function. If the former, presumably ponderosa pine has steadily taken up more water from near the soil surface and Douglas-fir has shifted uptake to a greater depth. If the latter, we suggest the pronounced changes in leaf-water delta(18)O are a result of changes in leaf structure and function with tree size and age.

Height growth determinants and adaptation to temperature in pines: a case study ofPinus contortaandPinus monticola

In this study we aimed to compare and explain the height growth performance of two contrasting pine species: lodgepole pine (Pinus contorta Dougl. ex. Loud) and western white pine(Pinus monticola Dougl. ex D. Don.). We compiled measurements of total height growth at different ages and shoot elongation phenology realized in several provenance test trials for 109 provenances of lodgepole pine and 54 provenances of western white pine. The response of shoot elongation to temperature was assessed using a phenological model fitted on provenance mean growth curves. Although total height growth followed the same geographic trends in both species, the response of shoot elongation to temperature was different between the two, with few (lodgepole pine) or no differences among provenances (western white pine) from diverse geographic regions. The temperature for which potential cell growth rate is 50% was 10.8 ± 0.13 °C (mean ± standard error) for western white pine compared to 5.26 ± 0.075 °C for lodgepole pine. Phenology did not explain growth performance differences among geographical regions in both species, which instead were explained by differences in the number of internodes set the preceding summer; provenances originating from stressful environments produced the fewest internodes, possibly due to reallocation of carbohydrates to stress resistance.

Exponential fertilization of Pinus monticola seedlings: nutrient uptake efficiency, leaching fractions, and early outplanting performance

We evaluated nutrient uptake efficiency and subsequent leaching fractions for western white pine (Pinus monticola Dougl. ex D. Don) seedlings grown with exponentially increasing or conventional (constant) fertilization in a greenhouse. Conventional fertilization was associated with higher leachate electrical conductivity and greater nutrient losses, which were more pronounced for the more mobile NO3-N than for NH4-N. Exponential fertilization increased seedling nitrogen (N) uptake efficiency (75%), which helped reduce leaching losses compared to conventional fertilization (50%). Although exponentially fertilized plants received 45% less fertilizer (20 mg N·plant–1) during the establishment and accelerated growth phases compared with conventional cohorts (36 mg N·plant–1), seedling morphological characteristics and nutrient status were similar at the end of greenhouse culture and after 2 years of growth in the field, except that exponential fertilization increased root volume after the first year and also increased ectomycorrhizal colonization. Reduction of applied fertilizer quantities and enhanced fertilizer uptake efficiency through exponential fertilization allows for production of high-quality seedlings while simultaneously minimizing fertilizer inputs and mitigating potential environmental contamination.

Water uptake and oil distribution during imbibition of seeds of western white pine (Pinus monticola Dougl. ex D. Don) monitored in vivo using magnetic resonance imaging

Dry or fully imbibed seeds of western white pine (Pinus monticola Dougl. ex D. Don) were studied using high-resolution magnetic resonance imaging (MRI). Analyses of the dry seed revealed many of the gross anatomical features of seed structure. Furthermore, the non-invasive nature of MRI allowed for a study of the dynamics of water and oil distribution during in situ imbibition of a single seed with time-lapse chemical shift selective MRI. During soaking of the dry seed, water penetrated through the seed coat and megagametophyte. The cotyledons of the embryo (located in the chalazal end of the seed) were the first to show hydration followed by the hypocotyl and later the radicle. After penetrating the seed coat, water in the micropylar end of the seed likely also contributed to further hydration of the embryo; however, the micropyle itself did not appear to be a site for water entry into the seed. A model that describes the kinetics of the earlier stages of imbibition is proposed. Non-viable pine seeds captured with MRI displayed atypical imbibition kinetics and were distinguished by their rapid and uncontrolled water uptake. The potential of MR microimaging for detailed studies of water uptake and distribution during the soaking, moist chilling ("stratification"), and germination of conifer seeds is discussed.

Characterization, expression and evolution of two novel subfamilies ofPinus monticolacDNAs encoding pathogenesis-related (PR)-10 proteins

Proteins of the pathogenesis-related (PR)-10 family are induced in many plants by phytopathogens and environmental stresses. A multi-gene family of PR10 proteins has previously been found in the genome of western white pine (Pinus monticola Dougl. ex D. Don). We isolated two novel subfamilies of PR10 cDNAs (PmPR10-2 and PmPR10-3) from P. monticola that are distinct from other PR10 genes (PmPR10-1.1-1.14) reported from the same species. The PmPR10 proteins are grouped in three subfamilies based on similarity in amino acid sequences. The sequence identities of PmPR10 proteins are much higher among members within a subfamily than among members of different subfamilies (86-99% versus 59-68%). Induction of both PmPR10-2 and PmPR10-3 mRNAs was detected by reverse transcriptase-polymerase chain reaction (RT-PCR) in needles in response to wounding treatment. PmPR10-3 was also expressed in needles during cold acclimation in winter. Transcript levels of both PmPR10-2 and PmPR10-3 were less than the detectable levels of constitutive expression in roots, stems and vegetative shoots, whereas PmPR10-1.10 mRNA of subfamily I was expressed at various levels. Phylogenetic analysis showed that PmPR10 and PR10 proteins from other conifers are grouped within one clade that is distinct from that of angiosperm PR10 proteins. In the conifer monophyletic group, PR10 sequences diversify into three distinct clusters. Among these three clusters, some PR10 proteins from single conifer species showed greater divergence distances than sequences from other conifer species, suggesting that, within the conifers, the multi-gene family underwent great diversification during evolution. Based on ratios of nonsynonymous to synonymous nucleotide substitutions (Ka/Ks), we speculate that positive selection resulted in the divergence of PmPR10 subfamilies I and III. Possible mechanisms and significance of PR10 gene evolution are discussed.

Western white pine growth relative to forest openings

In northern Rocky Mountains moist forests, timber harvesting, fire exclusion, and an introduced stem disease have contributed to the decline in western white pine (Pinus monticola Dougl. ex D. Don) abundance (from 90% to 10% of the area). Relations between canopy openings (0.1–15 ha) and western white pine growth within different physical settings are identified. Objectives include relating western white pine seedling and sapling growth to canopy opening attributes (defined by fisheye photography), identifying western white pine competitive thresholds (occupancy, competitive advantage, free-to-grow status) in relation to opening size, and relating canopy opening attributes to overstory density descriptors. We sampled 620 western white pine plus competing trees, canopy opening characteristics, landscape position, and overstory density descriptors. Analysis included log-linear and polynomial regression. Visible sky (canopy opening) and tree age were significantly related to growth of selected pines. Radiation explained less variation in growth than canopy opening. Thresholds for western white pine to occupy a site (&gt;23% canopy opening), gain a competitive advantage (&gt;50% canopy opening), and achieve free-to-grow status (&gt;92% canopy opening) over western hemlock were developed. There was a significant although weak (R2 &lt; 0.12) relation between canopy opening and overstory density descriptors.

Bedrock type significantly affects individual tree mortality for various conifers in the inland Northwest, U.S.A.

Individual tree mortality models for western white pine (Pinus monticola Dougl. ex D. Don), Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), grand fir (Abies grandis (Dougl. ex D. Don) Lindl.), western redcedar (Thuja plicata Donn ex. D. Don), western hemlock (Tsuga heterophylla (Raf.) Sarg.), and western larch (Larix occidentalis Nutt.) were developed using data from permanent research plots located throughout the inland Northwest. The proposed linear logistic models included the following independent variables: diameter at breast height, basal area in larger trees, basal area per hectare, individual tree diameter increment, and a set of dummy variables for the six bedrock types, which were granite, basalt, metasedimentary, sedimentary, mixed – glacial till, and deep deposit. The results show that rock type significantly affects individual tree mortality for western white pine, Douglas-fir, and western redcedar, while grand fir, western hemlock, and western larch were not affected. Western white pine and Douglas-fir growing on meta sedimentary rocks exhibited greater mortality probabilities than on other rocks. Mortality probabilities for western hemlock were low across all rock types, including "nutrient-poor" rocks like metasedimentary types.

Dormancy termination of western white pine ( Pinus monticola Dougl. Ex D. Don) seeds is associated with changes in abscisic acid metabolism

Western white pine (Pinus monticola) seeds exhibit deep dormancy at maturity and seed populations require several months of moist chilling to reach their uppermost germination capacities. Abscisic acid (ABA) and its metabolites, phaseic acid (PA), dihydrophaseic acid (DPA), 7'-hydroxy ABA (7'OH ABA) and ABA-glucose ester (ABA-GE), were quantified in western white pine seeds during dormancy breakage (moist chilling) and germination using an HPLC-tandem mass spectrometry method with multiple reaction monitoring and internal standards incorporating deuterium-labeled analogs. In the seed coat, ABA and metabolite levels were high in dry seeds, but declined precipitously during the pre-moist-chilling water soak to relatively low levels thereafter. In the embryo and megagametophyte, ABA levels decreased significantly during moist chilling, coincident with an increase in the germination capacity of seeds. ABA catabolism occurred via several routes, depending on the stage and the seed tissue. Moist chilling of seeds led to increases in PA and DPA levels in both the embryo and megagametophyte. Within the embryo, 7'OH ABA and ABA-GE also accumulated during moist chilling; however, 7'OH ABA peaked early in germination. Changes in ABA flux, i.e. shifts in the ratio between biosynthesis and catabolism, occurred at three distinct stages during the transition from dormant seed to seedling. During moist chilling, the relative rate of ABA catabolism exceeded ABA biosynthesis. This trend became even more pronounced during germination, and germination was also accompanied by a decrease in the ABA catabolites DPA and PA, presumably as a result of their further metabolism and/or leaching/transport. The transition from germination to post-germinative growth was accompanied by a shift toward ABA biosynthesis. Dormant imbibed seeds, kept in warm moist conditions for 30 days (after an initial 13 days of soaking), maintained high ABA levels, while the amounts of PA, 7'OH ABA, and DPA decreased or remained at steady-state levels. Thus, in the absence of conditions required to break dormancy there were no net changes in ABA biosynthesis and catabolism.