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The invasion of non-native plants can alter the diversity and activity of soil microorganisms and nutrient cycling within forests. We used field studies to analyze the impact of a successful invasive groundcover, Alliaria petiolata, on fungal diversity, soil nutrient availability, and pH in five northeastern US forests. We also used laboratory and greenhouse experiments to test three mechanisms by which A. petiolata may alter soil processes: (1) the release of volatile, cyanogenic glucosides from plant tissue; (2) the exudation of plant secondary compounds from roots; and (3) the decomposition of litter. Fungal community composition was significantly different between invaded and uninvaded soils at one site. Compared to uninvaded plots, plots invaded by A. petiolata were consistently and significantly higher in N, P, Ca and Mg availability, and soil pH. In the laboratory, the release of volatile compounds from the leaves of A. petiolata did not significantly alter soil N availability. Similarly, in the greenhouse, the colonization of native soils by A. petiolata roots did not alter soil nutrient cycling, implying that the exudation of secondary compounds has little effect on soil processes. In a leaf litter decomposition experiment, however, green rosette leaves of A. petiolata significantly increased the rate of decomposition of native tree species. The accelerated decomposition of leaf litter from native trees in the presence of A. petiolata rosette leaves shows that the death of these high-nutrient-content leaves stimulates decomposition to a greater extent than any negative effect that secondary compounds may have on the activity of the microbes decomposing the native litter. The results presented here, integrated with recent related studies, suggest that this invasive plant may change soil nutrient availability in such a way as to create a positive feedback between site occupancy and continued proliferation.

The responses of subtropical conifer plantation to climate change scenarios were investigated in Qianyanzhou by the process-based physiological-ecological model EALCO (ecological assimilation of land and climate observation). The results showed that CO2 concentration had the greatest effects on the carbon and water fluxes of the plantation, followed by temperature, and precipitation. CO2 concentration was the main driving factor for the gross photosynthesis productivity of this plantation ecosystem, and temperature and CO2 concentration were the key environmental factors controlling the ecosystem respiration. Increasing temperature accelerated the respiration of aboveground part dramatically, while increasing CO2 concentration had greater effects on soil respiration. The evapotranspiration was enhanced by increasing temperature, but reduced by increasing CO2 concentration. Under the future climate changing scenario (the year 2100), the net primary productivity of this plantation ecosystem would be increased by 22%, suggesting that this ecosystem is still capable of sequestrating carbon.

The occurrence and abundance of conifers along climate gradients in the Inland Northwest (USA) was assessed using data from 5082 field plots, 81% of which were forested. Analyses using the Random Forests classification tree revealed that the sequential distribution of species along an altitudinal gradient could be predicted with reasonable accuracy from a single climate variable, a growing-season dryness index, calculated from the ratio of degree-days >5 degrees C that accumulate in the frost-free season to the summer precipitation. While the appearance and departure of species in an ascending altitudinal sequence were closely related to the dryness index, the departure was most easily visualized in relation to negative degree-days (degree-days < 0 degrees C). The results were in close agreement with the works of descriptive ecologists. A Weibull response function was used to predict from climate variables the abundance and occurrence probabilities of each species, using binned data. The fit of the models was excellent, generally accounting for >90% of the variance among 100 classes.

Leaf mass per area (LMA) is one of the most widely measured of all plant functional traits. In deciduous forests, there is similarity between plastic and evolutionary responses of LMA to light gradients. In evergreens, however, LMA is lower in shaded than sunlit individuals of the same species, whereas shade-tolerant evergreens have higher LMA than light-demanders grown under the same conditions. We suggest that this pattern of 'counter-gradient variation' results from some combination of (i) close evolutionary coordination of LMA with leaf lifespan, (ii) selection for different leaf constitutions (relative investment in cell walls versus cell contents) in sun and shade environments and/or (iii) constraints on plasticity as a result of genetic correlations between phenotypes expressed in sun and shade.

The invasive Mediterranean pine engraver, Orthotomicus erosus (Wollaston) (Coleoptera: Scolytidae), was detected in North America in 2004, and it is currently distributed in the southern Central Valley of California. It originates from the Mediterranean region, the Middle East, and Asia, and it reproduces on pines (Pinus spp.). To identify potentially vulnerable native and adventive hosts in North America, no-choice host range tests were conducted in the laboratory on 22 conifer species. The beetle reproduced on four pines from its native Eurasian range--Aleppo, Canary Island, Italian stone, and Scots pines; 11 native North American pines--eastern white, grey, jack, Jeffrey, loblolly, Monterey, ponderosa, red, Sierra lodgepole, singleleaf pinyon, and sugar pines; and four native nonpines--Douglas-fir, black and white spruce, and tamarack. Among nonpines, fewer progeny developed and they were of smaller size on Douglas-fir and tamarack, but sex ratios of progeny were nearly 1:1 on all hosts. Last, beetles did not develop on white fir, incense cedar, and coast redwood. With loblolly pine, the first new adults emerged 42 d after parental females were introduced into host logs at temperatures of 20-33 degrees C and 523.5 or 334.7 accumulated degree-days based on lower development thresholds of 13.6 or 18 degrees C, respectively.

The boreal coniferous forests form the most extended vegetation zone of the Northern Hemisphere. As opposed to North America, they are disconnected from the mountain coniferous forests in Europe, because of the dominant east-west direction of the mountain chains. Consequently, the mountain forests show some unique characteristic features of glacial survival and postglacial history, as well. The mountain coniferous forests have numerous common floral and faunal elements with the boreal zone. However, the few unique faunal elements of the European mountain coniferous forests can be used to unravel the peculiar patterns and processes of this biome. In this issue of Molecular Ecology, Thomas Schmitt and Karola Haubrich (2008) use the relatively common and taxonomically well-studied butterfly, the large ringlet (Erebia euryale) to identify the last glacial refugia and postglacial expansion routes.

There has been considerable recent interest in how human-induced species loss affects community and ecosystem properties. These effects are particularly apparent when a commercially valuable species is harvested from an ecosystem, such as occurs through single-tree harvesting or selective logging of desired timber species in natural forests. In New Zealand mixed-species rain forests, single-tree harvesting of the emergent gymnosperm Dacrydium cupressinum, or rimu, has been widespread. This harvesting has been contentious in part because of possible ecological impacts of Dacrydium removal on the remainder of the forest, but many of these effects remain unexplored. We identified an area where an unintended 40-year "removal experiment" had been set up that involved selective extraction of individual Dacrydium trees. We measured aboveground and belowground variables at set distances from both individual live trees and stumps of trees harvested 40 years ago. Live trees had effects both above and below ground by affecting diversity and cover of several components of the vegetation (usually negatively), promoting soil C sequestration, enhancing ratios of soil C:P and N:P, and affecting community structure of soil microflora. These effects extended to 8 m from the tree base and were likely caused by poor-quality litter and humus produced by the trees. Measurements for the stumps revealed strong legacy effects of prior presence of trees on some properties (e.g., cover by understory herbs and ferns, soil C sequestration, soil C:P and N:P ratios), but not others (e.g., soil fungal biomass, soil N concentration). These results suggest that the legacy of prior presence of Dacrydium may remain for several decades or centuries, and certainly well over 40 years. They also demonstrate that, while large Dacrydium individuals (and their removal) may have important effects in their immediate proximity, within a forest, these effects should only be important in localized patches containing high densities of large trees. Finally, this study emphasizes that deliberate extraction of a particular tree species from a forest can exert influences both above and below ground if the removed species has a different functional role than that of the other plant species present.

By using LI-8100 CO2 Flux Meter, the CO2 flux of soil surface and the temperature and moisture content at soil depth of 10 cm in conifer/broad-leaved mixed forests under different intensity of selective cutting in Dailing Forest Bureau of Heilongjiang Province were determined simultaneously from 2003 to 2007, with the changes of soil surface CO2 flux after selective cutting and their affecting factors studied. The results showed that the CO2 flux was correlated with soil temperature and moisture content, with a relatively high confidence ranged from 68% to 98%. Selective cutting increased the CO2 flux, with an average annual increment being from 7.17% to 26.89%. There was a significant quadratic correlation between soil surface CO2 flux and cutting intensity (R2 = 0.961). Soil organic matter content and cutting intensity were the main factors affecting the changes of soil surface CO2 flux after selective cutting.

Bordered pits are cavities in the lignified cell walls of xylem conduits (vessels and tracheids) that are essential components in the water-transport system of higher plants. The pit membrane, which lies in the center of each pit, allows water to pass between xylem conduits but limits the spread of embolism and vascular pathogens in the xylem. Averaged across a wide range of species, pits account for > 50% of total xylem hydraulic resistance, indicating that they are an important factor in the overall hydraulic efficiency of plants. The structure of pits varies dramatically across species, with large differences evident in the porosity and thickness of pit membranes. Because greater porosity reduces hydraulic resistance but increases vulnerability to embolism, differences in pit structure are expected to correlate with trade-offs between efficiency and safety of water transport. However, trade-offs in hydraulic function are influenced both by pit-level differences in structure (e.g. average porosity of pit membranes) and by tissue-level changes in conduit allometry (average length, diameter) and the total surface area of pit membranes that connects vessels. In this review we address the impact of variation in pit structure on water transport in plants from the level of individual pits to the whole plant.

An assessment of the insect guild structure associated with immature and mature eastern hemlock, Tsuga canadensis (L.) Carrière, at high and low elevations was made before the invasion by the hemlock woolly adelgid, Adelges tsugae (Annand), and elongate hemlock scale, Fiorinia externa Ferris. Guild dynamics were determined among 243 insect species associated with eastern hemlock in the southern Appalachians. Detritivore, hematophage, herbivore, fungivore, parasitoid, predator, scavenger, and transient guild assignments were made. The herbivore and transient guilds were subdivided into chewers, sapsuckers, flower feeders, pollen feeders, pollen/nectar/sap feeders, and seed feeders. A significant association was found between guilds on immature and mature hemlocks at low elevations, but no significant association was found at high elevations. There was also a strong association of phytophagous insects on immature hemlocks and scavengers on mature hemlocks. The observed transient, scavenger, and predator guilds had a significantly lower co-occurrence among species indicating segregation of the species within these guilds. The herbivore guild had a significantly higher co-occurrence among species than the simulated index indicating aggregation of species within this guild across sites. Detritivore, hematophage, and parasitoid guilds did not differ significantly from the simulated community. These results document a diverse and dynamic insect community on eastern hemlock before invasion by hemlock woolly adelgid and elongate hemlock scale.

In many pine species (Family Pinaceae), ovulate cones structurally resemble a turbine, which has been widely interpreted as an adaptation for improving pollination by producing complex aerodynamic effects. We tested the turbine interpretation by quantifying patterns of pollen accumulation on ovulate cones in a wind tunnel and by using simulation models based on computational fluid dynamics. We used computer-aided design and computed tomography to create computational fluid dynamics model cones. We studied three species: Pinus radiata, Pinus sylvestris, and Cedrus libani. Irrespective of the approach or species studied, we found no evidence that turbine-like aerodynamics made a significant contribution to pollen accumulation, which instead occurred primarily by simple impaction. Consequently, we suggest alternative adaptive interpretations for the structure of ovulate cones.

The early genetic effects of habitat degradation were investigated in the critically endangered conifer Araucaria nemorosa. This species occurs in New Caledonia, a global biodiversity hotspot where the world's greatest concentration of endemic conifer species coincides with an extremely high level of habitat destruction due to fire and mining. Using seven microsatellite loci, estimates were made of genetic marker variation, inbreeding coefficients and population differentiation of adult and seedling cohorts of A. nemorosa. These were contrasted with equivalent estimates, made over similar spatial scales and with the same marker loci, in the locally common and more widespread sister species Araucaria columnaris. There were no significant differences in population genetic parameters between adult populations of the two species, despite their different abundances. However, in A. nemorosa, the juvenile cohort showed a loss of rare alleles and elevated levels of inbreeding when compared to the adult cohort. These genetic differences between the cohorts were not observed in the locally common A. columnaris. This suggests that recent environmental degradation is influencing the genetic structure of A. nemorosa populations. Although this is not detectable among predisturbance adult populations, an early warning of these impacts is evident in more recently established seedling cohorts. The conservation implications of these results are discussed.

Plant cold hardiness has complicated attributes. This paper introduced the research advances in establishing the dynamic models of coniferous trees cold hardiness, with the advantages and disadvantages of the models presented and the further studies suggested. In the models established initially, temperature was concerned as the only environmental factor affecting the cold hardiness, and the concept of stationary level of cold hardiness was introduced. Due to the obvious prediction errors of these models, the stationary level of cold hardiness was modeled later by assuming the existence of an additive effect of temperature and photoperiod on the increase of cold hardiness. Furthermore, the responses of the annual development phases for cold hardiness to environment were considered. The model researchers have paid more attention to the additive effect models, and run some experiments to test the additivity principle. However, the research results on Scots pine (Pinus sylvestris) indicated that its organs did not support the presumption of an additive response of cold hardiness by temperature and photoperiod, and the interaction between environmental factors should be taken into account. The mathematical models of cold hardiness need to be developed and improved.

Soil water potential (Psi(s)) is often estimated by measuring leaf water potential before dawn (Psi(pd)), based on the assumption that the plant water status has come into equilibrium with that of the soil. However, it has been documented for a number of plant species that stomata do not close completely at night, allowing for nocturnal transpiration and thus preventing nocturnal soil-plant water potential equilibration. The potential for nighttime transpiration necessitates testing the assumption of nocturnal equilibration before accepting Psi(pd) as a valid estimate of Psi(s). We determined the magnitude of disequilibrium between Psi(pd) and Psi(s) in four temperate conifer species across three height classes through a replicated study in northern Idaho. Based on both stomatal conductance and sap flux measurements, we confirmed that the combination of open stomata and high nocturnal atmospheric vapor pressure deficit (D) resulted in nocturnal transpiration in all four species. Nocturnal stomatal conductance (g(s-noc)) averaged about 33% of mid-morning conductance values. We used species-specific estimates of g(s-noc) and leaf specific conductance to correct Psi(pd) values for nocturnal transpiration at the time the samples were collected. Compared with the unadjusted values, corrected values reflected a significantly higher Psi(pd) (when D > 0.12 kPa). These results demonstrate that comparisons of Psi(pd) among species, canopy height classes and sites, and across growing seasons can be influenced by differential amounts of nocturnal transpiration, leading to flawed results. Consequently, it is important to account for the presence of nocturnal transpiration, either through a properly parameterized model or by making Psi(pd) measurements when D is sufficiently low that it cannot drive nocturnal transpiration. Violating these conditions will likely result in underestimation of Psi(s).

The study on the density of ambient particles settling upon the leaf surface of six conifers in Beijing, the micro-configurations of the leaf surface, and the mineral and element compositions of the particles showed that at the same sites and for the same tree species, the density of the particles settling upon leaf surface increased with increasing ambient pollution, but for various tree species, it differed significantly, with the sequence of Sabina chinensis and Platycladus orientalis > Cedrus deodara and Pinus bungeana > P. tabulaeformis and Picea koraiensis. Due to the effects of road dust, low height leaf had a larger density of particles. The density of the particles was smaller in summer than in winter because of the rainfall and new leaf growth. The larger the roughness of leaf surface, the larger density of the particles was. In the particles, the overall content of SiO2, CaCO3, CaMg(CO3,), NaCl, 2CaSO4 . H2O, CaSO4 . 2H2O and Fe2O3 was about 10%-30%, and the main minerals were montmorillonite, illite, kaolinite and feldspar. The total content of 21 test elements in the particles reached 16%-37%, among which, Ca, Al, Fe, Mg, K, Na and S occupied 97% or more, while the others were very few and less affected by sampling sites and tree species.

The gaseous environment surrounding parenchyma in woody tissue is low in O2 and high in CO2, but it is not known to what extent this affects respiration or might play a role in cell death during heartwood formation. Sapwood respiration was measured in two conifers and three angiosperms following equilibration to levels of O2 and CO2 common within stems, using both inner and outer sapwood to test for an effect of age. Across all species and tissue ages, lowering the O2 level from 10% to 5% (v/v) resulted in about a 25% decrease in respiration in the absence of CO2, but a non-significant decrease at 10% CO2. The inhibitory effect of 10% CO2 was smaller and only significant at 10% O2, where it reduced respiration by about 14%. Equilibration to a wider range of gas combinations in Pinus strobus L. showed the same effect: 10% CO2 inhibited respiration by about 15% at both 20% and 10% O2, but had no net effect at 5% O2. In an extreme treatment, 1% O2+20% CO2 increased respiration by over 30% relative to 1% O2 alone, suggesting a shift in metabolic response to high CO2 as O2 decreases. Although an increase in respiration would be detrimental under limiting O2, this extreme gas combination is unlikely to exist within most stems. Instead, moderate reductions in respiration under realistic O2 and CO2 levels suggest that within-stem gas composition does not severely limit respiration and is unlikely to cause the death of xylem parenchyma during heartwood formation.

This study presents the impacts of clear-cutting and site preparation on soil and needle 15N-fractionation of Scots pine (Pinus sylvestris, L.) and Norway spruce (Picea abies (L.), Karst). Three microsites on different methods of site preparation were used: (i) mound (broken O/E/B horizons piled upside down over undisturbed humus), (ii) deep (exposed C-horizon) and (iii) shallow (exposed E/B horizon). We found significant differences between species, between closed forest and clear-cuts as well as between different site preparations. For instance, in the context of interspecific variations, the mean needle nitrogen concentrations of both seedlings (1.15,+/-0.10 %) and mature (1.09,+/-0.07 %) pine trees were significantly higher compared to corresponding needle concentrations of seedlings (0.88,+/-0.06 %) and mature trees (0.79,+/-0.02 %) of spruce. Similarly, we observed significantly more 15N-enriched needles of mature spruces (-4.0,+/-0.20 per thousand) as well as of seedlings (-5.0,+/-0.11 per thousand) relative to that of mature pine needles (-5.6,+/-0.10 per thousand) and seedlings (-6.0,+/-0.31 per thousand). These variations were assumed to be caused by the variation in mycorrhizal associations between the species. We assume that the proportion of mycorrhizal N-uptake of pines might have been larger than that of spruce. Regarding the clear-cut effects on N and 15N of both tree species, we observed that, in the mature natural stand, needle N concentrations of both pine (1.09,+/-0.07 %) and spruce (0.79,+/-0.02 %) tree species did not change significantly after clear-cutting (pine: 1.01,+/-0.06 %; spruce: 0.74,+/-0.04 % ). However, clear-cutting resulted in the significant increase in needle 15N natural abundance of both pine (-2.70,+/-0.06 per thousand) and spruce (-2.09,+/-0.05 per thousand) in comparison to that of natural stand (pine:-5.60,+/-0.10 per thousand; spruce:-4.00,+/-0.20 per thousand), which is assumed to be due to the increased level of nitrification and leaching of nitrate after clear-cutting. In the context of site preparation methods, soil and needle N were observed to be more 15N-enriched in deep and shallow treatment sites compared to that of closed forest site and untreated clear-cut site, which indicated that the main source of N uptake seems to be mainly directly from the soil of the corresponding horizons of mineral soil with higher delta15N.

Theoretical and empirical studies have long suggested that stability and complexity are intimately related, but evidence from long-lived systems at large scales is lacking. Stability can either be driven by complex species interactions, or it can be driven by the presence/absence and abundance of a species best able to perform a specific ecosystem function. We use 64 years of stand productivity measures in forest systems composed of four dominant conifer tree species to contrast the effect of species richness and abundance on three stability measures. To perform this contrast, we measured the annual growth increments of > 900 trees in mixed and pure forest stands to test three hypotheses: increased species richness will (1) decrease stand variance, (2) increase stand resistance to drought events, and (3) increase stand resilience to drought events. In each case, the alternate hypothesis was that species richness had no effect, but that species composition and abundance within a stand drove variance, resistance, and resilience. In pure stands, the four species demonstrated significant differences in productivity, and in their resistance and resilience to drought events. The two pine species were the most drought resistant and resilient, whereas mountain hemlock was the least resistant and resilient, and red fir was intermediate. For community measures we found a moderately significant (P = 0.08) increase in the community coefficient of variation and a significant (P = 0.03) increase in resilience with increased species richness, but no significant relationship between species richness and community resistance, though the variance in community resistance to drought decreased with species richness. Community resistance to drought was significantly (P = 0.001) correlated to the relative abundance of lodgepole pine, the most resistant species. We propose that resistance is driven by competition for a single limiting resource, with negative diversity effects. In contrast resilience measures the capacity of communities to partition resources in the absence of a single limiting resource, demonstrating positive diversity effects.

Wood structure and function of juvenile wood from 18 conifer species from four conifer families (Araucariaceae, Cupressaceae, Pinaceae and Podocarpaceae) were examined for a trade-off between wood reinforcement and hydraulic efficiency. Wood density and tracheid 'thickness-to-span' ratio were used as anatomical proxies for mechanical properties. The thickness:span represented the ratio of tracheid double wall thickness to lumen diameter. Hydraulic resistivity (R) of tracheids on a cross-sectional area basis (RCA) increased over 50-fold with increasing density and thickness:span, implying a strength versus efficiency conflict. The conflict arose because density and thickness:span were increased by narrowing tracheid diameter rather than by thickening walls, which may be developmentally difficult. In the Pinaceae and Cupressaceae species, density and thickness:span correlated strongly with protection from drought-induced embolism, suggesting that mechanical strength was required in part to withstand tracheid collapse by negative sap pressure. These species showed a corresponding trade-off between increasing RCA and embolism protection. In contrast, species of Podocarpaceae and Araucariaceae were overbuilt for their embolism protection and were hydraulically inefficient, having greater density, thickness:span and RCA, none of which were correlated with vulnerability to embolism.

Fast-growing exotic trees are widely planted in the tropics to counteract deforestation; however, their patterns of water use could be detrimental to overall ecosystem productivity through their impact on ecosystem water budget. In a comparative field study on seasonal soil-plant water dynamics of two exotic species (Cupressus lusitanica Mill. and Eucalyptus globulus Labill.) and the indigenous Podocarpus falcatus (Thunb.) Mirb. in south Ethiopia, we combined a 2.5-year record for climate and soil water availability, natural-abundance oxygen isotope ratios (delta(18)O) of soil and xylem water, destructive root sampling and transpiration measurements. Soil was generally driest under C. lusitanica with its dense canopy and shallow root system, particularly following a relatively low-rainfall wet season, with the wettest soil under E. globulus. Wet season transpiration of C. lusitanica was twice that of the other species. In the dry season, P. falcatus and C. lusitanica reduced transpiration by a factor of six and two, respectively, whereas E. globulus showed a fivefold increase. In all species, there was a shift in water uptake to deeper soil layers as the dry season progressed, accompanied by relocation of live fine root biomass (LFR) of C. lusitanica and P. falcatus to deeper layers. Under P. falcatus, variability in soil matric potential, narrow delta(18)O depth gradients and high LFR indicated fast water redistribution. Subsoil water uptake was important only for E. globulus, which had low topsoil LFR and tap roots exploiting deep water. Although P. falcatus appeared better adapted to varying soil water availability than the exotic species, both conifers decreased growth substantially during dry weather. Growth of E. globulus was largely independent of topsoil water content, giving it the potential to cause substantial dry-season groundwater depletion.

Polar regions were covered with extensive forests during the Cretaceous and early Tertiary, and supported trees comparable in size and productivity to those of present-day temperate forests. With a winter of total or near darkness and a summer of continuous, low-angle illumination, these temperate, high-latitude forests were characterized by a light regime without a contemporary counterpart. Although maximum irradiances were much lower than at mid-latitudes, the 24-h photoperiod provided similar integrated light flux. Taxodium, Larix and Metasequoia, three genera of deciduous conifers that occurred in paleoarctic wet forests, have extant, closely related descendents. However, the contemporary relative abundance of these genera differs greatly from that in the paleoarctic. To provide insight into attributes that favor competitive success in a continuous-light environment, we subjected saplings of these genera to a natural photoperiod or a 24-h photoperiod and measured gas exchange, chlorophyll fluorescence, non-structural carbohydrate concentrations, biomass production and carbon allocation. Exposure to continuous light significantly decreased photosynthetic capacity and quantum efficiency of photosystem II in Taxodium and Larix, but had minimal influence in Metasequoia. In midsummer, foliar starch concentration substantially increased in both Taxodium and Larix saplings grown in continuous light, which may have contributed to end-product down-regulation of photosynthetic capacity. In contrast, Metasequoia allocated photosynthate to continuous production of new foliar biomass. This difference in carbon allocation may have provided Metasequoia with a two fold advantage in the paleoarctic by minimizing depression of photosynthetic capacity and increasing photosynthetic surface.

Changes in the 2H and 18O of atmospheric water vapour provide information for integrating aspects of gas exchange within forest canopies. In this study, we show that diurnal fluctuations in the oxygen isotope ratio (delta 18O) as high as 4% per hundred were observed for water vapour (delta (18)Ovp) above and within an old-growth coniferous forest in the Pacific Northwest region of the United States. Values of delta 18Ovp decreased in the morning, reached a minimum at midday, and recovered to early-morning values in the late afternoon, creating a nearly symmetrical diurnal pattern for two consecutive summer days. A mass balance budget was derived and assessed for the 18O of canopy water vapour over a 2-d period by considering the 18O-isoflux of canopy transpiration, soil evaporation and the air entering the canopy column. The budget was used to address two questions: (1) do delta 18O values of canopy water vapour reflect the biospheric influence, or are such signals swamped by atmospheric mixing? and (2) what mechanisms drive temporal variations of delta 18Ovp? Model calculations show that the entry of air into the canopy column resulted in an isotopically depleted 18O-isoflux in the morning of day 1, causing values of delta 18Ovp, to decrease. An isotopically enriched 18O-isoflux resulting from transpiration then offset this decreased delta 18Ovp later during the day. Contributions of 18O-isoflux from soil evaporation were relatively small on day 1 but were more significant on day 2, despite the small H2(16)O fluxes. From measurements of leaf water volume and sapflux, we determined the turnover time of leaf water in the needles of Douglas-fir trees as approximately 11 h at midday. Such an extended turnover time suggests that transpiration may not have occurred at the commonly assumed isotopic steady state. We tested a non-steady state model for predicting delta 18O of leaf water. Our model calculations show that assuming isotopic steady state increased isoflux of transpiration. The impact of this increase on the modelled delta 18Ovp was clearly detectable, suggesting the importance of considering isotopic non-steady state of transpiration in studies of forest 18O water balance.

Conifer foliage structures affect branch and bud temperature by altering the development and convective resistance of the thermal boundary layer. This paper examines foliage effects on forced convection in branches and buds of Picea glauca (Moench) Voss and Pinus contorta Dougl. Ex. Loud., two species that represent the range of variation in foliage structure among conifers. Forced convection is characterized by a power law relating Nusselt (heat transfer) and Reynolds (boundary layer development) numbers. Data were collected in a laminar flow wind tunnel for free stream velocities of 0.16-6.95 m s(-1). Scaling parameters were compared against literature values for silver cast branch replicas, a bed of real foliage, cylinders, and tube banks. Foliage structures reduced Nusselt numbers (heat transfer) relative to cylinders, which are typically used to approximate leafless branches and buds. Significantly different scaling relationships were observed for all foliage structures considered. Forced convection scaling relationships varied with foliage structure. The scaling relationships reported here account for variation within populations of branches and buds and can be used to characterize forced convection in a forest canopy.

Intra-annual radial growth rates and durations in trees are reported to differ greatly in relation to species, site and environmental conditions. However, very similar dynamics of cambial activity and wood formation are observed in temperate and boreal zones. Here, we compared weekly xylem cell production and variation in stem circumference in the main northern hemisphere conifer species (genera Picea, Pinus, Abies and Larix) from 1996 to 2003. Dynamics of radial growth were modeled with a Gompertz function, defining the upper asymptote (A), x-axis placement (beta) and rate of change (kappa). A strong linear relationship was found between the constants beta and kappa for both types of analysis. The slope of the linear regression, which corresponds to the time at which maximum growth rate occurred, appeared to converge towards the summer solstice. The maximum growth rate occurred around the time of maximum day length, and not during the warmest period of the year as previously suggested. The achievements of photoperiod could act as a growth constraint or a limit after which the rate of tree-ring formation tends to decrease, thus allowing plants to safely complete secondary cell wall lignification before winter.

Insects select their hosts, but trees cannot select which herbivores will feed upon them. Thus, as long-lived stationary organisms, conifers must resist the onslaught of varying and multiple attackers over their lifetime. Arguably, the greatest threats to conifers are herbivorous insects and their associated pathogens. Insects such as bark beetles, stem- and wood-boring insects, shoot-feeding weevils, and foliage-feeding budworms and sawflies are among the most devastating pests of conifer forests. Conifer trees produce a great diversity of compounds, such as an enormous array of terpenoids and phenolics, that may impart resistance to a variety of herbivores and microorganisms. Insects have evolved to specialize in resistance to these chemicals -- choosing, feeding upon, and colonizing hosts they perceive to be best suited to reproduction. This review focuses on the plant-insect interactions mediated by conifer-produced terpenoids. To understand the role of terpenoids in conifer-insect interactions, we must understand how conifers produce the wide diversity of terpenoids, as well as understand how these specific compounds affect insect behaviour and physiology. This review examines what chemicals are produced, the genes and proteins involved in their biosynthesis, how they work, and how they are regulated. It also examines how insects and their associated pathogens interact with, elicit, and are affected by conifer-produced terpenoids.

Throughfall and open field bulk precipitation were measured at three coniferous sites during 1995-2002 in the framework of ICP Integrated Monitoring and at five coniferous sites during 1996-2002 in the framework of ICP Forests (Level II). The coniferous canopies acted as a sink for nitrate and ammonium and as a source for base cations: Ca(2+), Mg(2+) and K(+). The estimated share of SO(4)-S dry deposition from total deposition was 1.5-4 times higher for dormant period compared to growing period. During the study period average annual throughfall and bulk deposition of SO(4)-S decreased significantly, 2.8 and 2.3 times, respectively. Throughfall enrichment with base cations increased in the order Mg < Na < Ca < K. Using Na as a tracer ion, average dry deposition and canopy leaching were calculated. Leaching was the dominant process for TF enrichment by potassium. Leaching of base cations occurred during growing as well as dormant period. The calculated internal flux of Ca(2+) and Mg(2+) varied in the range of 0.6-2.0 and 0.6-1.2 kg ha(-1) per year in spruce and pine stands, respectively. The internal circulation of K(+) was significantly higher (8.9-10.9 kg ha(-1) per year) in spruce stands than in pine stands (2.7-4.4 kg ha(-1) per year).

The unicellular conifer tracheid should have greater flow resistance per length (resistivity) than the multicellular angiosperm vessel, because its high-resistance end-walls are closer together. However, tracheids and vessels had comparable resistivities for the same diameter, despite tracheids being over 10 times shorter. End-wall pits of tracheids averaged 59 times lower flow resistance on an area basis than vessel pits, owing to the unique torus-margo structure of the conifer pit membrane. The evolution of this membrane was as hydraulically important as that of vessels. Without their specialized pits, conifers would have 38 times the flow resistance, making conifer-dominated ecosystems improbable in an angiosperm world.

Ecosystem process models are often used to predict carbon flux on a landscape or on a global scale. Such models must be aggregate and canopies are often treated as a uniform unit of foliage. Parameters that are known to vary within the canopy, e.g., nitrogen content and leaf mass per area, are often estimated by a mean value for the canopy. Estimating appropriate means is complicated, especially in mixed-species stands and in complex terrain. We analyzed sources of variation in specific parameters with the goal of testing various simplifying assumptions. The measurements came from mixed-species forests in the northern Rocky Mountains. We found that, for three important parameters (nitrogen concentration and content, and leaf mass per area), a sample taken near the vertical center of the crown provided a good estimate of the mean values for the crown. Altitude (700-1700 m), solar insolation (4200-5400 MJ m(-2) year(-1)) and leaf area index (1-11) had negligible effects on the parameters; only species differences were consistently detected. The correlation between mass-based photosynthetic rates and mass-based nitrogen concentrations was much weaker than the correlation between area-based photosynthetic rates and area-based nitrogen concentration. Comparison of photosynthesis-nitrogen relationships for a wide variety of conifer species and sites revealed a broad general trend that can be used in models. These results suggest important potential simplifications in model parameterization, most notably that canopy means can be estimated with ease, that complex terrain is a minor source of variation in these parameters and that use of one photosynthesis-nitrogen relationship for conifer species does not result in large errors. Species-to-species variation, however, was large and needs to be accounted for when parameterizing process models.

Few studies have examined the effects of plant growth on nutrient remobilization in phenologically contrasting species. Here we evaluated the consequences of above-ground seasonality of growth and leaf shedding on the remobilization of nutrients from branches in eight evergreen Mediterranean phanaerophytes that differ widely in phenology. Vegetative growth, flower bud formation, flowering, fruiting, leaf shedding, and the variations in nitrogen (N), phosphorus (P) and potassium (K) pools in branches throughout the year were monitored in each species. Nitrogen and P remobilization occurred in summer, after vegetative growth and synchronously with leaf shedding. Despite the time-lag between growth and remobilization, the branches that invested more nutrients in vegetative growth also remobilized more nutrients from their old organs. Potassium remobilization peaked in the climatically harshest periods, and appears to be related to osmotic requirements. We conclude that N and P remobilization occurs mainly associated with leaf senescence, which might be triggered by factors such as the replenishment of nutrient reserves in woody organs, the hormonal relations between new and old leaves, or the constraints that summer drought poses on the amount of leaf area per branch in summer.

Conifers are long-lived organisms, and part of their success is due to their potent defense mechanisms. This review focuses on bark defenses, a front line against organisms trying to reach the nutrient-rich phloem. A major breach of the bark can lead to tree death, as evidenced by the millions of trees killed every year by specialized bark-invading insects. Different defense strategies have arisen in conifer lineages, but the general strategy is one of overlapping constitutive mechanical and chemical defenses overlaid with the capacity to up-regulate additional defenses. The defense strategy incorporates a graded response from 'repel', through 'defend' and 'kill', to 'compartmentalize', depending upon the advance of the invading organism. Using a combination of toxic and polymer chemistry, anatomical structures and their placement, and inducible defenses, conifers have evolved bark defense mechanisms that work against a variety of pests. However, these can be overcome by strategies including aggregation pheromones of bark beetles and introduction of virulent phytopathogens. The defense structures and chemicals in conifer bark are reviewed and questions about their coevolution with bark beetles are discussed., (Copyright New Phytologist (2005).)

We assessed the relative limitations to photosynthesis imposed by stomatal and non-stomatal processes in Dacrydium cupressinum Lamb. (Podocarpaceae), which is the dominant species in a native, mixed conifer-broad-leaved rainforest in New Zealand. For comparison, we included three co-occurring broad-leaved tree species (Meterosideros umbellata Cav. (Myrtaceae), Weinmannia racemosa L.f. (Cunoniaceae) and Quintinia acutifolia Kirk (Escalloniaceae)) that differ in phylogeny and in leaf morphology from D. cupressinum. We found that low foliage phosphorus content on an area basis (P(a)) limited light-saturated photosynthesis on an area basis (A(sat)) in Q. acutifolia. Depth in the canopy did not generally affect A(sat) or the relative limitations to A(sat) because of stomatal and non-stomatal constraints, despite reductions in the ratio of foliage mass to area, foliar nitrogen on an area basis (N(a)) and P(a) with depth in the canopy. In the canopy-dominant conifer D. cupressinum, A(sat) was low, consistent with low values of the maximum rate of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) carboxylation (V(cmax)). In comparison, the A(sat) response of the three broad-leaved tree species was quite variable. Although A(sat) was high in the canopy-dominant M. umbellata, it was low in the sub-canopy trees W. racemosa and Q. acutifolia. Relative stomatal limitation to photosynthesis was more pronounced in W. racemosa (40%) than in the other three species (28-33%). Despite differences in degree, non-stomatal limitation to A(sat) predominated in all tree species.

* The hydraulic plumbing of vascular plant leaves varies considerably between major plant groups both in the spatial organization of veins, as well as their anatomical structure. * Five conifers, three ferns and 12 angiosperm trees were selected from tropical and temperate forests to investigate whether the profound differences in foliar morphology of these groups lead to correspondingly profound differences in leaf hydraulic efficiency. * We found that angiosperm leaves spanned a range of leaf hydraulic conductance from 3.9 to 36 mmol m2 s-1 MPa-1, whereas ferns (5.9-11.4 mmol m-2 s-1 MPa-1) and conifers (1.6-9.0 mmol m-2 s-1 MPa-1) were uniformly less conductive to liquid water. Leaf hydraulic conductance (Kleaf) correlated strongly with stomatal conductance indicating an internal leaf-level regulation of liquid and vapour conductances. Photosynthetic capacity also increased with Kleaf, however, it became saturated at values of Kleaf over 20 mmol m-2 s-1 MPa-1. * The data suggest that vessels in the leaves of the angiosperms studied provide them with the flexibility to produce highly conductive leaves with correspondingly high photosynthetic capacities relative to tracheid-bearing species.

Just as a soggy paper straw is prone to yielding under the applied suction of a thirsty drinker, the xylem tracheids in leaves seem prone to collapse as water potential declines, impeding their function. Here we describe the collapse, under tension, of lignified cells peripheral to the leaf vein of a broad-leaved rainforest conifer, Podocarpus grayi de Laub. Leaves of Podocarpus are characterized by an array of cylindrical tracheids aligned perpendicular to the leaf vein, apparently involved in the distribution of water radially through the mesophyll. During leaf desiccation the majority of these tracheids collapsed from circular to flat over the water potential range -1.5 to -2.8 MPa. An increase in the percentage of tracheids collapsed during imposed water stress was mirrored by declining leaf hydraulic conductivity (K(leaf)), implying a direct effect on water transport efficiency. Stomata responded to water stress by closing at -2.0 MPa when 45% of cells were collapsed and K(leaf) had declined by 25%. This was still substantially before the initial indications of cavitation-induced loss of hydraulic conductance in the leaf vein, at -3 MPa. Plants droughted until 49% of tracheids had collapsed were found to fully recover tracheid shape and leaf function 1 week after rewatering. A simple mechanical model of tracheid collapse, derived from the theoretical buckling pressure for pipes, accurately predicted the collapse dynamics observed in P. grayi, substantiating estimates of cell wall elasticity and measured leaf water potential. The possible adaptive advantages of collapsible vascular tissue are discussed.

The structure and dynamics of coniferous forests in Changbai Mountain were studied at different spatial scales, including ground survey of permanent plots and analysis of multitemporal satellite images. Plot-scale examinations showed that the mortality rate was 7% - 9%, and the recruitment rate was 18% - 20% per 10 years. Species composition changed over time. Picea jezoensis var. microsperma, Abies nephrolepis and Betula ermanii presented a self-maintaining capability, because they could regenerate under canopy. Larix olgensis was a pioneer species and could regenerate only in open land or gaps. This species played an important role by providing conditions for the regeneration of spruce and fir. The tree density in the mature forest was 1 000 stems x hm(-2) for trees bigger than 3 cm in diameter, which showed no significant variations among different stands. Landsat TM images were used for detecting the cover changes from 1984 to 1997. Large scales of wind throw were detected by this approach. Based on t he analysis of radiance changes at the landscape scale, the pixel number of the disturbed area was similar to that of the succeeding stands, suggesting that the forest was in a state of equilibrium. Fine gaps, however, were difficult to identify with the TM data because of its coarse resolution. The mosaic structure ofthe subalpine vegetation was characterized by scattered larch patches. At the landscape level, the vegetation was in a stable stage.

Conifers have evolved constitutive and inducible defense mechanisms to help in both wound healing and defense against attack by bark beetles and other organisms. These defenses include oleoresin, phenolics, and static structures in secondary phloem, such as lignified cells and calcium oxalate crystals, that create physical barriers. We used a phylogenetic approach to investigate the defense anatomy of conifer stems of 13 species from five families following treatment with methyl jasmonate (MJ), a compound that induces defense responses in stems of several Pinaceae species. Methyl jasmonate induced a response similar to wounding except that the response was not accompanied by lesion formation, necrosis or a hypersensitive response. In the Pinaceae species studied, MJ induced polyphenolic parenchyma (PP) cell activation and xylem traumatic resin duct (TD) formation. Members of the Taxodiaceae, which are not known to produce large quantities of resin, showed massive xylem TD formation and surface resinosis following MJ treatment. Treatment with MJ caused members of the Araucariaceae and Cupressaceae to form axial phloem resin ducts but not xylem ducts, whereas Podocarpaceae species showed no induction of resin-producing structures. All species treated with MJ showed phenolic deposition in PP cells, and early lignification of phloem fibers was observed in most of the non-Pinaceae species. We conclude that, although evolution of resin-producing structures occurred independently in conifer lineages, MJ seems to induce resin production regardless of tissue location, as well as inducing deposition of phenolic compounds. Co-evolution of conifer defensive strategies and bark beetle pests is discussed.

Survival after freezing was measured for seeds and germinants of four seedlots each of interior spruce (Picea glauca x engelmannii complex), lodgepole pine (Pinus contorta Dougl. ex Loud.), Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) and western red cedar (Thuja plicata Donn ex D. Donn). Effects of eight seed treatments on post-freezing survival of seeds and germinants were tested: dry, imbibed and stratified seed, and seed placed in a growth chamber for 2, 5, 10, 15, 20 or 30 days in a 16-h photoperiod and a 22/17 degrees C thermoperiod. Survival was related to the water content of seeds and germinants, germination rate and seedlot origin. After freezing for 3 h at -196 degrees C, dry seed of most seedlots of interior spruce, Douglas-fir and western red cedar had 84-96% germination, whereas lodgepole pine seedlots had 53-82% germination. Freezing tolerance declined significantly after imbibition in lodgepole pine, Douglas-fir and interior spruce seed (western red cedar was not tested), and mean LT50 of imbibed seed of these species was -30, -24.5 and -20 degrees C, respectively. Freezing tolerance continued to decline to a minimum LT50 of -4 to -7 degrees C after 10 days in a growth chamber for interior spruce, Douglas-fir and lodgepole pine, or after 15 days for western red cedar. Minimum freezing tolerance was reached at the stage of rapid hypocotyl elongation. In all species, a slight increase in freezing tolerance of germinants was observed once cotyledons emerged from the seed coat. The decrease in freezing tolerance during the transition from dry to germinating seed correlated with increases in seed water content. Changes in freezing tolerance between 10 and 30 days in the growth chamber were not correlated with seedling water content. Within a species, seedlots differed significantly in freezing tolerance after 2 or 5 days in the growth chamber. Because all seedlots of interior spruce and lodgepole pine germinated quickly, there was no correlation between seedlot hardiness and rate of germination. Germination rate and freezing tolerance of Douglas-fir and western red cedar seedlots was negatively correlated. There was a significant correlation between LT50 after 10 days in the growth chamber and minimum spring temperature at the location of seedlot origin for interior spruce and three seedlots of western red cedar, but no relationship was apparent for lodgepole pine and Douglas-fir.

Deciduous forests covered the ice-free polar regions 280 to 40 million years ago under warm "greenhouse" climates and high atmospheric pCO2. Their deciduous habit is frequently interpreted as an adaptation for minimizing carbon losses during winter, but experiments with "living fossils" in a simulated warm polar environment refute this explanation. Measured carbon losses through leaf abscission of deciduous trees are significantly greater than losses through winter respiration in evergreens, yet annual rates of primary productivity are similar in all species. Here, we investigate mechanisms underlying this apparent paradox by measuring the seasonal patterns of leaf photosynthesis (A) under pCO2 enrichment in the same trees. During spring, A increased significantly in coastal redwood (Sequoia sempervirens), dawn redwood (Metasequoia glyptostroboides), and swamp cypress (Taxodium distichum) at an elevated pCO2 of 80 Pa compared with controls at 40 Pa. However, strong acclimation in Rubisco carboxylation capacity (Vc,max) completely offset the CO2 response of A in all species by the end of 6 weeks of continuous illumination in the simulated polar summer. Further measurements demonstrated the temporary nature of acclimation, with increases in Vc,max during autumn restoring the CO2 sensitivity of A. Contrary to expectations, the acclimation of Vc,max was not always accompanied by accumulation of leaf carbohydrates, but was associated with a decline in leaf nitrogen in summer, suggesting an alteration of the balance in plant sources and sinks for carbon and nitrogen. Preliminary calculations using A indicated that winter carbon losses through deciduous leaf abscission and respiration were recovered by 10 to 25 d of canopy carbon fixation during summer, thereby explaining the productivity paradox.

We tested the hypotheses that freezing-induced embolism is related to conduit diameter, and that conifers and angiosperms with conduits of equivalent diameter will exhibit similar losses of hydraulic conductivity in response to freezing. We surveyed the freeze-thaw response of conifers with a broad range of tracheid diameters by subjecting wood segments (root, stem and trunk wood) to a freeze-thaw cycle at -0.5 MPa in a centrifuge. Embolism increased as mean tracheid diameter exceeded 30 microm. Tracheids with a critical diameter greater than 43 microm were calculated to embolize in response to freezing and thawing at a xylem pressure of -0.5 MPa. To confirm that freezing-induced embolism is a function of conduit air content, we air-saturated stems of Abies lasiocarpa (Hook.) Nutt. (mean conduit diameter 13.7 +/- 0.7 microm) by pressurizing them 1 to 60 times above atmospheric pressure, prior to freezing and thawing. The air saturation method simulated the effect of increased tracheid size because the degree of super-saturation is proportional to a tracheid volume holding an equivalent amount of dissolved air at ambient pressure. Embolism increased when the dissolved air content was equivalent to a mean tracheid diameter of 30 microm at ambient air pressure. Our centrifuge and air-saturation data show that conifers are as vulnerable to freeze-thaw embolism as angiosperms with equal conduit diameter. We suggest that the hydraulic conductivity of conifer wood is maximized by increasing tracheid diameters in locations where freezing is rare. Conversely, the narrowing of tracheid diameters protects against freezing-induced embolism in cold climates.

Freezing and thawing lead to xylem embolism when gas bubbles caused by ice formation expand during the thaw process. However, previous experimental studies indicated that conifers are resistant to freezing-induced embolism, unless xylem pressure becomes very negative during the freezing. In this study, we show that conifers experienced freezing-induced embolism when exposed to repeated freeze-thaw cycles and simultaneously to drought. Simulating conditions at the alpine timberline (128 days with freeze-thaw events and thawing rates of up to 9.5 K h(-1) in the xylem of exposed twigs during winter), young trees of Norway spruce [Picea abies (L.) Karst.] and stone pine (Pinus cembra L.) were exposed to 50 and 100 freeze-thaw cycles. This treatment caused a significant increase in embolism rates in drought-stressed samples. Upon 100 freeze-thaw cycles, vulnerability thresholds (50% loss of conductivity) were shifted 1.8 MPa (Norway spruce) and 0.8 MPa (stone pine) towards less negative water potentials. The results demonstrate that freeze-thaw cycles are a possible reason for winter-embolism in conifers observed in several field studies. Freezing-induced embolism may contribute to the altitudinal limits of conifers.

Annual or biannual time courses of plant delta13C (delta13C(p)) over the last century (70-100 years) were recorded for leafbases of four grasstrees (Xanthorrhoea preissii) at four sites in mediterranean Australia and wood of four conifers (Widdringtonia cedarbergensis) at two sites in mediterranean South Africa. There was a strong downward trend of 2-5.5(per thousand ) from 1935 to 1940 to the present in the eight plants. Trends were more variable from 1900 to 1940 with plants at two sites of each species showing an upward trend of 1-2.5 per thousand. Accepting that delta13C of the air (delta13C(a)) fell by almost 2 per thousand over the last century, the ratio of leaf intercellular CO2 to atmospheric CO2 (c(i)/c(a)) rose in five plants and remained unchanged in three over that period. Changes in c(i)/c(a) rather than delta13C(a) were more closely correlated with changes in delta13C(p) and accounted for 6.7-71.8% (22.6 c(i)/c(a)) and 28.2-93.3% (delta13C(a)) of the variation in delta13C(p). We doubt that possible changing patterns of rainfall, water availability, temperature, shade, air pollution or clearing for agriculture have contributed to the overall trend for c(i)/c(a) to rise over time. Instead, we provide evidence (concentrations of Fe and Mn in the grasstree leafbases) that decreasing photosynthetic capacity associated with falling nutrient availability due to the reduced occurrence of fire may have contributed to rising c(i)/c(a). Intrinsic water-use efficiency (W(i)) as a function of (c(a)-c(i)) usually increased linearly over the period, with the two exceptions explained by their marked increase in c(i)/c(a). We conclude that grasstrees may provide equivalent delta13C(p )and W(i) data to long-lived conifers and that their interpretation requires a consideration of the causes of variation in both c(i)/c(a )and delta13C(a).

Genetic variation in branchlet nutrient (N, P, K, Na, Ca, Mg, Mn and Fe) concentrations and mineral concentration (sum of branchlet P, K, Na, Ca, Mg, Mn and Fe concentrations) of 8-9-year-old hoop pine (Araucaria cunninghamii Ait. ex D. Don) half-sib families was assessed for four canopy positions at a wet site (23 families) and two canopy positions at an N- and water-limiting dry site (22 families) in relation to tree growth and associated branchlet carbon (delta13C) and oxygen (delta18O) isotope composition in southeast Queensland, Australia. Branchlet nutrient and mineral concentrations varied significantly among families and with canopy position and site. Depending on the canopy position sampled, the hoop pine family effect accounted for 0 to 13.8% of the total variation in branchlet N concentration, and for 0 to 30.3% of the total variation in branchlet mineral concentration at the wet site. The corresponding values for the family effect at the dry site were 0-13.3% for branchlet N concentration and 0-25.7% for branchlet mineral concentration. There were significant variations in branchlet P, K, Ca and Mg concentrations at both sites, and these variations differed with canopy position. Relationships between family means of branchlet N concentration and tree growth or delta13C or delta18O varied with canopy position at both sites. At the wet site, there were significant positive correlations between branchlet mineral concentration in the upper-outer or upper-inner canopy and tree height (r = 0.26 and 0.37, P < 0.01) and between branchlet mineral concentration and delta13C (r = 0.24, P < 0.01) in the upper-inner canopy, and a significant negative correlation between branchlet mineral concentration and delta13C (r = -0.21, P < 0.05) in the upper-outer canopy. At the dry site, branchlet mineral concentrations in the upper-inner and upper-outer canopy were significantly correlated with branchlet delta13C (r = -0.28 and -0.51, P < 0.01), and branchlet N concentration in the upper-inner canopy was significantly correlated with tree growth (r = 0.29, P < 0.01). A significant correlation between branchlet delta18O (an index of stomatal conductance) and branchlet mineral concentration at the dry site (r = 0.39, P = 0.020) indicated that stomatal conductance might be a factor regulating the variation in branchlet mineral concentration of the hoop pine families. Both branchlet N concentration and mineral concentration at particular canopy positions assist in selecting hoop pine families with improved tree growth and N- and water-use efficiency in environments where both N deficiency and a limited water supply are major factors affecting plantation productivity.

Boundaries between different forest types in Changbai Mountain Eastern China are results from complex interactions between forest ecosystems, topography, and geomorphology. Detecting and quantifying the transitional zones are highly important since high environmental heterogeneity and biodiversity are often found within these zones. In this study, we used GIS and multivariate statistics techniques (PCA and MSWA) to analyze data from Landsat TM satellite imageries and quantitatively determined the positions and widths of the landscape boundary between mountain birch and evergreen coniferous forests in the northern slope of Changbai Mountain. The results showed that the widths of the landscape boundary ranges from 30-50 m while using the MSWA or/and PC method. Such detected widths are consistent with field transect data that suggests a 50 m transitional zone width. The results further suggest that TM data can be used in combination with GIS and statistical techniques in determining forest landscape boundaries; MSWA is more reliable than PCA, while PCA can also be used to determine the landscape boundary when transects are properly located.

The storage, biomass, and decomposition of fallen woods in dark coniferous forest on the north slope of Changbai Mountain at elevations of 1260 m and 1620 m were studied. The results showed that the volume of fallen woods was 180.87 m3.hm-2, occupying 21.83% of the stand volume. The volume and biomass of fallen woods at elevation of 1260 m were higher than those at 1620 m. The volume of fallen woods was 52.57 m3.hm-2 at 1260 m, and was 193.85 m3.hm-2 at 1620 m, and their biomass was 6.21 t.hm-2 at 1260 m, and was 53.33 t.hm-2 at 1620 m. Monomial exponential attenuation model could be used to simulate the decomposition process of fallen woods. The results indicated that the decomposition constants of fallen woods were different among different tree species and different elevations. The decomposition constant of fallen woods of Abies was higher than that of Picea, and was higher at 1260 m than at 1620 m. The results also showed that the decomposition of Picea lasted longer.

We investigated effects of stem phloem girdles on apical control of branch angle, stem and branch growth and stem air content in six conifer species. A stem girdle 2 cm above a branch caused the branch to bend upward in all six species. Upward bending was associated with increased formation and action of compression wood (CW) in the lower portion of the branch. Compression wood also formed in the main stem below the branch, suggesting increased auxin production in the branch. A stem girdle 2 cm below a branch (the branch remained directly connected to the apex and distal branches) released the branch from apical control in Tsuga canadensis (L.) Carr., Pinus contorta Dougl. ex Loud. and Pseudotsuga menziesii (Mirb.) Franco. The branch bent up, but there was no CW formation in the stem. In Pinus rigida Mill., the branch exhibited increased cambial activity but did not bend up. A stem girdle > 20 cm below a branch did not release the branch from apical control in any of the species. These results support the hypothesis that branches compete with the subjacent stem for branch-produced photosynthate and that when the branch lacks this competitive sink it is released from apical control. A stem girdle 2 cm below a branch did not cause release of apical control in either Juniperus virginiana L. or Picea abies (L.) Karst. In these species, decreased shoot elongation and cambial activity above the girdle probably prevented release. A stem girdle 2 cm below a branch increased air content in the stem below the girdle in four of five species, whereas the other girdle treatments had no significant effect on stem air content. Although growth was inhibited above the girdle in the two species with the largest increase in air content, growth was not inhibited in the other species. High air content in stem segments isolated from distal auxin and carbohydrate sources is consistent with the hypothesis that a carbohydrate supply is required to refill embolized cells.