Vesicular mycorrhizal colonization of seedlings of Pinaceae and Betulaceae after spore inoculation with Glomus intraradices

 Although Pinaceae and Betulaceae have been reported to contain Glomus–type root endophytes, its ecological importance and the conditions influencing this symbiosis are poorly understood. Seedlings of Abies lasiocarpa, Alnus rubra, Pinus contorta, Pinus ponderosa, Pseudotsuga menziesii, and Tsuga heterophylla were inoculated with Glomus intraradices to determine the vesicular-arbuscular mycorrhizae (VAM) development and responsiveness of these hosts. The role of companion VAM host plants on mycorrhizal colonization and nutrient uptake by Pseudotsuga menziesii was also examined by growing seedlings of Pseudotsuga menziesii in dual culture with VAM hosts Thuja plicata or Calamagrostis rubescens. After 8 weeks, no seedlings were colonized. At 16 weeks, 8 of 17 Thuja plicata seedlings grown with Pseudotsuga menziesii and all 18 inoculated Thuja plicata seedlings grown alone were colonized with vesicles and hyphae. Two of 17 inoculated Pseudotsuga menziesii seedlings grown in dual culture with Thuja plicata were colonized with abundant vesicles and hyphae. No ectomycorrhizal seedlings grown in monoculture were colonized. At 9 months, all 10 Calamagrostis rubescens and all 10 inoculated Pseudotsuga menziesii seedlings grown in dual culture were colonized by vesicles and hyphae. Two of 10 inoculated Pseudotsuga menziesiiand 1 of 10 inoculated Pinus ponderosa seedlings grown in monoculture were similarly colonized. The mean phosphorus content in the needles of colonized Pseudotsuga menziesii seedlings grown with Calamagrostis rubescens was about twice as high as in noncolonized Pseudotsuga menziesiiseedlings grown with Calamagrostis rubescens. Tissue nitrogen did not differ between these treatments. The results show that Glomus intraradices colonization of Pinaceae is most successful when a VAM host is present, although some vesicular colonization of Pinaceae occurred in the absence of a VAM host. Accepted: 24 September 1997     

Late quaternary dynamics of pollen influx at mineral lake,Washington

Pollen influx analysis at Mineral Lake, Washington, indicates that immediately south of the Puget Lobe of the Fraser Glaciation, tundra was a characteristic vegetation until 16,300 years ago. Invasion ofPinus contorta began 17,500 years B.P., and boreal climax conifers (Abies, Picea andTsuga mertensiana), 16,300, but was temporarily interrupted by the Vashon advance (14,500–14,000 yr B.P.).Pseudotsuga menziesii began to grow in population 10,750 years ago, and woodland was established within a time span of 1,000 years. Modern lowland coniferous forests began to form 7,000 years ago. Logistic analysis of pollen abundance changes show that the intrinsic growth rate,r (yr⁻¹), of pioneer species (e.g. 0.024–0.026 inPteridium aquilinum) is higher than that of climax species (e.g. 0.003 inThuja plicata).P. menziesii, a subclimax species, shows an intermediater value (0.013) between these two ecologically different taxa. The absoluter value ofP. contorta (−0.011) is only slightly lower than that ofP. menziesii, although their replacement began almost simultaneously. Thus competition between these species was intense before the inflection point ofPinus curve 10,100 years ago. At this time, forest gaps became abundantly available forPseudotsuga, as indicated by a peak of the diagnostic factor (the reciprocal of the pollen influx).     

The Importance of Large-Diameter Trees to Forest Structural Heterogeneity

Large-diameter trees dominate the structure, dynamics and function of many temperate and tropical forests. However, their attendant contributions to forest heterogeneity are rarely addressed. We established the Wind River Forest Dynamics Plot, a 25.6 ha permanent plot within which we tagged and mapped all 30,973 woody stems ≥1 cm dbh, all 1,966 snags ≥10 cm dbh, and all shrub patches ≥2 m². Basal area of the 26 woody species was 62.18 m²/ha, of which 61.60 m²/ha was trees and 0.58 m²/ha was tall shrubs. Large-diameter trees (≥100 cm dbh) comprised 1.5% of stems, 31.8% of basal area, and 17.6% of the heterogeneity of basal area, with basal area dominated by Tsuga heterophylla and Pseudotsuga menziesii. Small-diameter subpopulations of Pseudotsuga menziesii, Tsuga heterophylla and Thuja plicata, as well as all tree species combined, exhibited significant aggregation relative to the null model of complete spatial randomness (CSR) up to 9 m (P≤0.001). Patterns of large-diameter trees were either not different from CSR (Tsuga heterophylla), or exhibited slight aggregation (Pseudotsuga menziesii and Thuja plicata). Significant spatial repulsion between large-diameter and small-diameter Tsuga heterophylla suggests that large-diameter Tsuga heterophylla function as organizers of tree demography over decadal timescales through competitive interactions. Comparison among two forest dynamics plots suggests that forest structural diversity responds to intermediate-scale environmental heterogeneity and disturbances, similar to hypotheses about patterns of species richness, and richness- ecosystem function. Large mapped plots with detailed within-plot environmental spatial covariates will be required to test these hypotheses.     

Anaerobic induced ethanol synthesis in the stems of greenhouse-grown conifer seedlings

Ethanol synthesis was induced in stem segments from greenhouse-grown conifer seedlings by placing them in a N₂ atmosphere at 30 °C for 24 h. Stems from ponderosa pine,Pinus ponderosa Dougl. ex Laws., sugar pine,Pinus lambertiana Dougl., Pacific silver fir,Abies amabalis Dougl. ex Forbes, and lodgepole pine,Pinus contorta Dougl. ex Loud, produced the highest quantities of ethanol. This group also had the smallest and slowest growing stems. Within each of these species the amount of ethanol produced was inversely related to the stem volume. Stems from western hemlock,Tsuga heterophylla (Raf.) Sarg., grand fir, Abies grandis Dougl. ex Forbes, Douglas-fir,Pseudotsuga menziesii (Mirb.) Franco, and western redcedar,Thuja plicata Donn ex D. Don, all produced equivalent but low ethanol concentrations. These species had the largest and fastest growing stems. In this group only grand fir exhibited an inverse relationship between ethanol concentrations and stem volume. The relative amounts of ethanol synthesized by stems from Douglas-fir, western hemlock and western redcedar seedlings were not the same as subsequently observed in logs from mature trees of the same species under field conditions. Differences in the anaerobic environments for the two stem types could have affected the quantities of ethanol produced. The observed high amounts of ethanol produced by the stems from pine species were discussed in terms of their ability to handle periods of anaerobic stress or hypoxia.     

Predicted short-term radial-growth changes of trees based on past climate on Vancouver Island,British Columbia

Tree-ring radial expansion estimator (TREE) is an integrated radial growth model that allows users to define short-term climate change scenarios to anticipate the impact upon mature trees found growing at high elevation on Vancouver Island, British Columbia. Five individualistic models were built to represent the radial growth behaviour of mountain hemlock (Tsuga mertensiana (Bong.) Carr), yellow-cedar (Chamaecyparis nootkatensis (D. Don) Spach), western red-cedar (Thuja plicata Donn), Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), and western hemlock (Tsuga heterophylla (Raf.) Sarg.) trees. The models were developed on climate-radial growth relationships incorporating Nanaimo climate station data, and were able to explain from 55 to 68 per cent of the variance in radial growth. The models can be run with modifications to yearly precipitation and temperature variables, giving the user the ability to investigate the radial-growth impacts of a wide range of possible climate change scenarios. Results from eight such scenarios show that species growing within their ecological limits illustrate a limited change in radial growth to forecasted climate, while species growing at an ecotonal boundary are usually very sensitive to a specific climate variables (e.g., July temperature). A forecasted alteration to this key variable will then radically alter the radial-growth rate of the species.     

A chalcone synthase/stilbene synthase DNA probe for conifers

A probe for chalcone synthase (CHS) was generated by PCR using chalcone synthase conserved sequences. The cloned PCR product has high similarity to both chalcone synthase and stilbene synthase sequences. The probe was used to examine the organization of chalcone synthase and stilbene synthase genes in Abies procera, Pinus lambertiana, P. monticola, Picea glauca, P. sitchensis, Pseudostuga menziesii, Taxus brevifolia, and Thuja plicata. A large number of hybridizing bands were found in all species except T. plicata which did not cross hybridize. The hybridization patterns are highly polymorphic between the species and are also polymorphic within several of them.     

Acetylene reduction in conifer logs during early stages of decomposition

Acetylene reduction was examined periodically for as long as 68 months in the outer and inner bark, sapwood, and heartwood of decaying logs of western hemlock [Tsuga heterophylla (Raf.) Sarg.] western redcedar (Thuja plicata D. Don), Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco], and Pacific silver fir (Abies amabilis Dougl. ex Forbes) in the western Oregon Cascade Mountains. Tissues from freshly cut logs from sound trees were unable to reduce acetylene. However, after 18 months of decomposition, acetylene reduction was found in all log tissues except heartwood. Over the 68-month study period, no significant relationship between reduction rate and tissue moisture was found. Acetylene reduction rates differed significantly among tissues, log species, and time of exposure to decomposers. Although acetylene reduction generally showed a steady increase with time, tissues of some species showed a more complex, nonlinear pattern of change. Although the amount of nitrogen fixed is low compared to the total present in decaying logs, it might be an important source of readily available nitrogen for the microbiota responsible for decomposition.Paper 2587, Forest Research Laboratory, Oregon State University, Corvallis.Paper 2587, Forest Research Laboratory, Oregon State University, Corvallis.     

Stem respiratory potential in six softwood and four hardwood tree species in the central cascades of Oregon

Mature and old growth trees of varying sapwood thickness were compared with regard to stem respiration. An increment core-based, laboratory method under controlled temperature was used to measure tissue-level respiration (termed respiratory potential) of ten different tree species. Bark (dead outer and live inner combined), sapwood, and heartwood thickness measurements were used to predict sapwood volume from stem diameter (including bark) for four of the ten species. These predictions of sapwood volume were used to scale respiratory potential to the main-bole level (excluding all branches). On the core level, species that maintained narrow sapwood (8–16% of bole radius) such as Pseudotusga menziesii, Taxus brevifolia, and Thuja plicata, had sapwood respiratory potentials in the lower bole that were 50% higher (P<0.05) than species with wide sapwood (>16% of bole radius), such as Abies amabilis, Pinus monticola, and Tsuga heterophylla. This pattern was not observed for inner bark respiratory potential, or for sapwood respiratory potential within the crown. On the main-bole level, respiratory potential per unit volume was inversely correlated to the live bole volumetric fraction (inner bark plus sapwood divided by whole bole volume) (Adj. R²=0.6). Specifically, tree species with 18–20% of the main bole alive potentially respired 1.3–3 times more per unit live bole volume than species with over 40%, suggesting that the live bole was less metabolically active in tree species that maintained large volumes of sapwood.     

Mycorrhiza-like interaction by Morchella with species of the Pinaceae in pure culture synthesis

 Isolates from two species of Morchella were tested for ability to form mycorrhizae in pure culture synthesis with Arbutus menziesii, Larix occidentalis, Pinus contorta, Pinus ponderosa, andPseudotsuga menziesii. Ectomycorrhizal structures (mantle and Hartig net) formed with the four species of the Pinaceae but not with A. menziesii. Results are compared to previous studies on morel mycorrhizae and discussed in an ecological context. Accepted: 23 October 1999     

Canopy Carbon Gain and Water Use: Analysis of Old-growth Conifers in the Pacific Northwest

This report summarizes our current knowledge of leaf-level physiological processes that regulate carbon gain and water loss of the dominant tree species in an old-growth forest at the Wind River Canopy Crane Research Facility. Analysis includes measurements of photosynthesis, respiration, stomatal conductance, water potential, stable carbon isotope values, and biogenic hydrocarbon emissions from Douglas-fir (Pseudotsuga menziesii), western hemlock (Tsuga heterophylla), and western red cedar (Thuja plicata). Leaf-level information is used to scale fluxes up to the canopy to estimate gross primary production using a physiology-based process model. Both light-saturated and in situ photosynthesis exhibit pronounced vertical gradients through the canopy, but are consistently highest in Douglas-fir, intermediate in western hemlock, and lowest in western red cedar. Net photosynthesis and stomatal conductance are strongly dependent on vapor-pressure deficit in Douglas-fir, and decline through the course of a seasonal drought. Foliar respiration is similar for Douglas-fir and western hemlock, and lowest for western red cedar. Water-use efficiency varied with species and tree height, as indexed using stable carbon isotopes values for foliage. Leaf water potential is most negative for Douglas-fir and similar for western hemlock and western red cedar. Terpene fluxes from foliage equal approximately 1% of the net carbon loss from the forest. Modeled estimates based on physiological measurements show gross primary productivity (GPP) to be about 22 Mg C m^–2 y^–1. Physiological studies will be necessary to further refine estimates of stand-level carbon balance and to make long-term predictions of changes in carbon balance due to changes in forest structure, species composition, and climate.     

Terpene production and growth of three Pacific Northwest conifers in response to simulated browse and nutrient availability

Natural variation in ungulate browsing behavior interferes with the understanding of plant morphological and biochemical responses to herbivory. To investigate mechanisms for recovery from herbivory, we examined growth patterns and biosynthesis of terpenoids under simulated browse (three clipping intensities) and supplemental mineral nutrition (four levels of controlled-release fertilization) for Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco], western hemlock (Tsuga heterophylla Raf. Sarg.), and western red-cedar (Thuja plicata Donn ex D. Don) seedlings on a reforestation site in Northwestern Oregon, USA, that was fenced to exclude ungulates. Higher clipping intensities increased relative height growth (at cost of diameter growth) for all the species. Only western red-cedar showed a decline in monoterpene concentrations with increasing clipping severity, suggesting prioritization in biosynthesis of terpenoids for this species. Douglas-fir and western hemlock responded to fertilization mostly through increased growth. Western red-cedar growth responses to fertilization were less pronounced, but monoterpene concentrations were 2–3 times higher compared to non-fertilized trees. Douglas-fir and western hemlock browse recovery and responses to fertilization consisted primarily of increased growth, while western red-cedar balanced growth promotion with production of chemical defense compounds. Our data suggests the evolution of species-dependent resource allocation strategies in response to both browse and soil nutrient availability.     

Does nitrogen availability control rates of litter decomposition in forests?

The effects of increased exogenous N availability on rates of litter decomposition were assessed in several field fertilization trials. In a jack pine (Pinus banksiana Lamb.) forest, needle litter decomposed at the same rate in control plots and in plots fertilized with urea and ammonium nitrate (1350 kg N ha^-1) with or without P and K. Mixed needle litter of western hemlock (Tsuga heterophylla (Raf.) Sarg.), western red cedar (Thuja plicata Donn) and Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) incubated in plots recently amended with sewage sludge (500 kg N ha^-1) lost less weight during 3 years than did litter in control plots. Forest floor material also decomposed more slowly in plots amended with sewage sludge. Paper birch (Betula papyrifera Marsh.) leaf litter placed on sewage sludge (1000 kg N ha^-1), pulp sludge, or sewage-pulp sludge mixtures decomposed at the same rate as leaf litter in control plots. These experiments demonstrate little effect of exogenous N availability on rates of litter decomposition.The influence of endogenous N availability on rates of litter decomposition was examined in a microcosm experiment. Lodgepole pine (Pinus contorta var. latifolia Engelm.) needle litter collected from N-fertilized trees (525 kg N ha^-1 in ammonium nitrate) were 5 times richer in N than needles from control trees (1.56% N versus 0.33% N in control trees), but decomposed at the same rate. Green needles from fertilized trees contained twice as much N as needles from control trees (1.91% N versus 0.88% N), but decomposed at the same rate. These experiments suggest that N availability alone, either exogenous or endogenous, does not control rates of litter decomposition. Increased N availability, through fertilization or deposition, in the absence of changes in vegetation composition, will not alter rates of litter decomposition in forests.     

Ordination of epiphytic bryophyte communities in a wet-temperate coniferous forest,South-Coastal British Columbia

Three coniferous tree species (Pseudotsuga menziesii, Tsuga heterophylla, and Thuja plicata) at Capilano Canyon, British Columbia, were studied for their epiphytic communities. Quantitative data were obtained for fourteen bryophyte species by sampling at heights of 0.5 m, 1 m and 2 m above ground level. Scapania bolanderi is an abundant and widely distributed species; Dicranum fuscescens, Bazzania denudata, Lepidozia reptans and Hypnum circinale are also common. An environmentally uniform study area was chosen to minimize the effects of factors other than bark and elevation on the distribution of epiphytes. In general, tree species are similar in terms of their epiphytic assemblages, but ordination methods using quantitative data exposed compositional variation that may be explained by differences in microclimate and bark-type. In this respect, the results point the way to further studies to examine epiphyte associations in closer detail, and to relate these associations to specific factors in the microenvironment.Nomenclature for vascular plants follows Hitchcock & Cronquist (1973); bryophyte nomenclature follows Schofield (1976) for mosses and Stotler & Crandall-Stotler (1977) for hepatics.We thank Dr. W. B. Schofield for aiding in bryophyte identification.     

Water balance of conifer logs in early stages of decomposition

Seasonal and long-term changes in the water balance of conifer logs during the first 8 years of decomposition were studied in an old-growth Pseudotsuga/Tsuga forest in the Oregon Cascade Mountains. Measurements were made of the moisture content of outer bark, inner bark, sapwood, and heartwood and of the flow of water into and out of logs of four species (Abies amabilis, Pseudotsuga menziesii, Thuja plicata, and Tsuga heterophylla). After the logs had decomposed from 1 to 2 years, 38–47% of the canopy throughfall landing upon them ran off the surface, 29–34% leached from the bottom, and 21–30% was absorbed and evaporated. After 8 years of decomposition, water entering and then leaching from logs increased 1.3 times while runoff decreased a similar amount. The proportion of water stored by and evaporated from logs in this study indicates that in old growth forests they may intercept 2–5% of the canopy throughfall to the forest floor and that, even in early stages of decomposition, they may affect the hydrological cycle of Pacific Northwest old-growth forests.This is paper 2945 of the Forest Research Laboratory, Oregon State University, Corvallis     

Availability and uptake of inorganic nitrogen in a mixed old-growth coniferous forest

Old-growth forest stands of mixed species composition provide the opportunity to study species-specific influences on soil properties. We monitored rates of nitrogen mineralization, nitrification and an index of ammonium and nitrate uptake in a mixed old-growth stand of Douglas-fir (Pseudotsuga menziesii), western hemlock (Tsuga heterophylla) and western redcedar (Thuja plicata) over a two-year period. Litter and mineral soil (0–10-cm depth) were sampled adjacent to ten large trees of each species. After initial characterization of litter and soil, buried bags were incubated in both layers for ca. 2-month intervals. Soil and litter pH was lowest near western hemlocks. Nitrification, nitrate concentrations, and percent uptake as nitrate differed among the tree species; rates were highest near western redcedars. For all species, percent nitrification and nitrate uptake rate were higher in soil than in litter. The results indicate species-specific effects on ammonium and nitrate production and uptake within this forest type.The research described in this article has been funded in part by the US Environmental Protection Agency. This document has been prepared at the EPA Environmental Research Laboratory in Corvallis, Oregon, through Contract No. 68-C8-0006 to ManTech Environmental Technology, Inc. It has been subjected to the Agency's peer and administrative review and approved for publication. Mention of trade names or commercial products does not constitute endorsement or recommendation for use.The research described in this article has been funded in part by the US Environmental Protection Agency. This document has been prepared at the EPA Environmental Research Laboratory in Corvallis, Oregon, through Contract No. 68-C8-0006 to ManTech Environmental Technology, Inc. It has been subjected to the Agency's peer and administrative review and approved for publication. Mention of trade names or commercial products does not constitute endorsement or recommendation for use.     

The population biology of Bromus tectorum in forests: effect of disturbance,grazing, and litter on seedling establishment and reproduction

Summary The effect of tree canopy, understory, herbivores, and litter depth on seedling establishment, survival, and reproduction of the alien grass, Bromus tectorum (cheatgrass), was examined in a series of experiments in four forest habitat types in western North America. Higher recruitment, survival, and reproduction on clearcuts, which would be expected if the overstory alone is limiting the distribution of cheatgrass in forests, were not observed. Removing the understory in an otherwise undisturbed Pinus ponderosa forest did, however, increase the emergence of B. tectorum, but plants in these experimentally-created openings were more vulnerable to grazing by small mammals. In contrast, removing the sparse understory in an Abies forest neither enhanced recruitment nor increased the incidence of grazing of B. tectorum seedlings. Regardless of the forest habitat, most grazed plants died before maturity; even fewer grazed plants produced seeds. Litter depth influenced both recruitment and biomass production: both the rate of germination and the size of resultant seedlings were lower on thick litter (6 cm) compared to results on thin litter (1.5 cm). In the more open Pinus ponderosa and Pseudotsuga menziesii forests, cheatgrass colonization may often occur in openings in the understory alone. Colonization in the more shady A. grandis and Thuja plicata forests is unlikely, however, unless the opening extends through both the understory and the overstory. As a result, cheatgrass is unlikely to increase in any of these forests unless the scale and incidence of disturbance increases substantially.     

Das Photosynthesevermögen einiger Immergrüner während des Winters und seine Reaktivierungsgeschwindigkeit nach scharfen Frösten

Das Photosynthesevermögen von Pinus cembra und Rhododendron ferrugineum sowie von Pinus contorta, Picea glauca und Pseudotsuga menziesii wurden kurz nach Entnahme der Zweigproben und nach Verwöhnung wiederholt gemessen. Erstere Pflanzen stammten vom Patscherkofel bei Innsbruck, 2 000 ü. NN und wurden bei 16 °C Lufttemperatur und 30 000 Lux gemessen; die letzteren drei Baumarten von Kananaskis, 1 400 m ü. NN, Alberta, Kanada, bei Bedingungen von 15 °C und 5 500 Lux. Tiefe Temperaturen reduzieren bei allen fünf Arten das Photosynthesevermögen ganz erheblich. Nach scharfen Frösten tritt bei allen Arten trotz optimaler Umweltsbedingungen im Licht kurz nach Entnahme Respiration auf. Die Reaktivierungsgeschwindigkeit des Photosyntheseapparates ist aber je nach Art verschieden. Pinus cembra und Pinus contorta sind erst nach längerer Zeit aus ihrer tiefen Winterruhe zu aktivieren (3 bzw. 10 Tage bis zu 50% CO₂-Aufnahme der Hochleistungsperiode). Rhododendron ferrugineum, Picea glauca und Pseudotsuga menziesii können zu allen Meßterminen nach weniger als eintägiger Verwöhnung CO₂ binden. Nach längerer Verwöhnung können sie alle — auch im Winter — sommerliche Photosynthesewerte erreichen. Die verschiedene Reaktion von Pinus cembra und Rhododendron ferrugineum läßt sich mit dem verschiedenen Resistenzverhalten und den unterschiedlichen Wuchsorten deuten. Das verschiedene Verhalten der drei Baumarten der Rocky Mountains weist keine Parallelen mit der Temperaturresistenz oder Knospenöffnung auf. Umfärbung, Wassergehalt und Chlorophyllaktivierung geben Anhaltspunkte zur Klärung der unterschiedlichen Reaktivierungsgeschwindigkeiten des Photosyntheseapparates nach Frösten. Warmwetterperioden während des Winters können daher von Pinus contorta zur Stoffproduktion nicht genutzt werden, während dies bei Picea und Pseudotsuga, sofern Bodenwasser verfügbar ist, möglich wäre. Diese Ergebnisse könnten eine unter vielen Ursachen für die natürliche Vegetationsfolge sein     

Defense mechanisms of conifers : relationship of monoterpene cyclase activity to anatomical specialization and oleoresin monoterpene content

Cell-free extracts from Pinus ponderosa Lawson (ponderosa pine) and Pinus sylvestris L. (Scotch pine) wood exhibited high levels of monoterpene synthase (cyclase) activity, whereas bark extracts of these species contained no detectable activity, and they inhibited cyclase activity when added to extracts from wood, unless polyvinylpyrrolidone was included in the preparation. The molecular mass of the polyvinylpyrrolidone added was of little consequence; however, polyvinylpolypyrrolidone (a cross-linked insoluble form of the polymer) was ineffective in protecting enzyme activity. Based on these observations, methods were developed for the efficient extraction and assay of monoterpene cyclase activity from conifer stem (wood and bark) tissue. The level of monoterpene cyclase activity for a given conifer species was shown to correlate closely with the monoterpene content of the oleoresin and with the degree of anatomical complexity of the specialized resin-secreting structures. Cyclase activity and monoterpene content were lowest in the stems of species containing only isolated resin cells, such as western red cedar (Thuja plicata D. Don). Increasing levels of cyclase activity and oleoresin monoterpenes were observed in advancing from species with multicellular resin blisters (true firs [Abies]) to those with organized resin passages, such as western larch (Larix occidentalis Nutt.), Colorado blue spruce (Picea pungens Engelm.) and Douglas-fir (Pseudotsuga menziesii [Mirb.] Franco). The highest levels of cyclase activity and oleoresin monoterpenes were noted in Pinus species that contain the most highly developed resin duct systems. The relationship between biosynthetic capacity, as measured by cyclase activity, monoterpene content, and the degree of organization of the secretory structures for a given species, may reflect the total number of specialized resin-producing cells per unit mass of stem tissue.     

Rates of stomatal opening in conifer seedlings in relation to air temperature and daily carbon gain

Experiments were conducted on well watered 1-year-old Douglas fir [Pseudotsuga menziesii (Mirb.) Franco], western hemlock [Tsuga heterophylla (Raf.) Sarg.] and western redcedar (Thuja plicata Donn) seedlings to determine the effects of temperature on whole-plant photosynthetic and stomatal responses to short-term fluctuations in irradiance (Q). Following a step change in Q, time constants (τ, the period over which 63% of the total change occurs) for stomatal conductance (g_s) and assimilation rate (A) decreased linearly with increasing air temperature (T_air). For example, in western redcedar τ_A decreased from 30 ± 4 min at 5 °C to 10 ± 1 min at 25 °C. In all cases, τ_A was within 10–15% of τ_gs. There was considerable variation in τ among individuals within a given species. Differences between species became more pronounced with decreasing temperature. Multiplicative models that included functions for τ accounted for 99% of the diurnal variability in A and g_s for seedlings exposed to varying air temperature, irradiance and vapour pressure deficit. Estimates of daily A were within 2% of those measured. Intermittent cloud cover and understory shading were approximated by exposing seedlings to 3–4 episodes (≥1 h) of shade (200 or 500 μmol m^?2 s^?1) or complete darkness during the day. In such cases, daily A was overestimated by up to 4 and 21%, respectively, if a function for τ was excluded from the models. Our results suggest that there is scope for selecting seedling stock for increased carbon assimilation on the basis of reduced time constants. For example, in western redcedar, a 40% reduction in τ could lead to increases in daily carbon gains of almost 5% depending on the frequency and degree of shading. If these daily gains were translated into increased dry matter production and compounded, seasonal gains would be even larger.     

The influence of canopy cover on understorey development in forests of the western Cascade Range,Oregon, USA

Natural disturbances, especially fire and treefalls, influence tree canopy composition in the Pseudotsuga menziesii forests of the western Cascade Range, Oregon. The composition of tree, shrub, and herb assemblages in the understorey of stands with different canopy types, such as maturing Pseudotsuga, Tsuga heterophylla, or mixed species stands, also differs.Differences in both canopy type and the prevalence of canopy openings correlated with different degrees of understorey development in stands of similar ages. This suggests that understorey assemblages also reflect disturbance history. Before understorey assemblages can be used to relate community samples to community or habitat types, the extent to which their composition reflects long term influences of stand history vs. differences in site potential must be determined.