Sustaining the growth of Pinaceae trees under nutrient-limited edaphic conditions via plant-beneficial bacteria.

Lodgepole pine, a prominent Pinaceae tree species native to western North America, is well-known for its ability to thrive in highly disturbed and degraded areas. One such area is the Sub-Boreal Pine-Spruce xeric-cold (SBPSxc) region in British Columbia, Canada, which is characterized by weakly-developed, parched soils that lack an organic forest floor and essential plant-available nutrients. We hypothesized that plant growth-promoting bacteria could play a significant role in sustaining the growth of lodgepole pine trees in the SBPSxc region. Testing this hypothesis, we evaluated plant growth-promoting abilities of six endophytic bacterial strains previously isolated from lodgepole pine trees growing in this region. These bacterial strains significantly enhanced the length and biomass of their natural host (lodgepole pine) as well as a foreign host (hybrid white spruce) in a 540-day long greenhouse trial. This growth stimulation could be linked to the diverse plant growth-promoting (PGP) abilities detected in these strains using in vitro assays for inorganic/organic phosphate-solubilization, siderophore production IAA production, ACC deaminase activity, lytic enzymes (chitinase, ß-1,3-glucanase, protease, and cellulase) activity, ammonia production and catalase activity. ACC deaminase activity was also detected in vivo for all strains using ethylene-sensitive plants-canola and tomato. Notably, strains belonging to the Burkholderiaceae family (HP-S1r, LP-R1r and LP-R2r) showed the greatest potential in all PGP assays and enhanced pine and spruce seedling length and biomass by up to 1.5-fold and 4-fold, respectively. Therefore, such bacterial strains with multifarious PGP abilities could be crucial for survival and growth of lodgepole pine trees in the SBPSxc region and could potentially be utilized as bioinoculant for Pinaceae trees in highly disturbed and nutrient-poor ecosystems.

Inter- and intra-tree variability of carbon and oxygen stable isotope ratios of modern pollen from nine European tree species.

Stable carbon and oxygen isotope ratios of raw pollen sampled from nine abundant tree species growing in natural habitats of central and northern Europe were investigated to understand the intra- and inter-specific variability of pollen-isotope values. All species yielded specific δ13Cpollen and δ18Opollen values and patterns, which can be ascribed to their physiology and habitat preferences. Broad-leaved trees flowering early in the year before leaf proliferation (Alnus glutinosa and Corylus avellana) exhibited on average 2.6‰ lower δ13Cpollen and 3.1‰ lower δ18Opollen values than broad-leaved and coniferous trees flowering during mid and late spring (Acer pseudoplatanus, Betula pendula, Carpinus betulus, Fagus sylvatica, Picea abies, Pinus sylvestris and Quercus robur). Mean species-specific δ13Cpollen values did not change markedly over time, whereas δ18Opollen values of two consecutive years were often statistically distinct. An intra-annual analysis of B. pendula and P. sylvestris pollen revealed increasing δ18Opollen values during the final weeks of pollen development. However, the δ13Cpollen values remained consistent throughout the pollen-maturation process. Detailed intra-individual analysis yielded circumferential and height-dependent variations within carbon and oxygen pollen-isotopes and the sampling position on a tree accounted for differences of up to 3.5‰ for δ13Cpollen and 2.1‰ for δ18Opollen. A comparison of isotope ranges from different geographic settings revealed gradients between maritime and continental as well as between high and low altitudinal study sites. The results of stepwise regression analysis demonstrated, that carbon and oxygen pollen-isotopes also reflect local non-climate environmental conditions. A detailed understanding of isotope patterns and ranges in modern pollen is necessary to enhance the accuracy of palaeoclimate investigations on δ13C and δ18O of fossil pollen. Furthermore, pollen-isotope values are species-specific and the analysis of species growing during different phenophases may be valuable for palaeoweather reconstructions of different seasons.

Intra-specific variation in growth and wood density traits under water-limited conditions: Long-term-, short-term-, and sudden responses of four conifer tree species.

Consequences of climate change will severely affect forest ecosystems in the near future, yet our understanding of how and why trees are responding to their abiotic environment is still limited. Intra-specific variation (ITV) in the growth response of trees to warming and drought has been widely neglected so far, but could play a key role for adapting forests to future climate conditions. We analyzed tree rings from four conifers (Picea abies, Abies alba, Larix decidua, Pseudotsuga menziesii) regarding their intra-specific adaptation potential when trees are growing at the warm and dry margins of species distributions. Our study comprises data from four common garden experiments (45 provenances and a total of 743 trees) and assessed growth response at different temporal scales from decades (long-term) to only a few event years (short-term) and finally for density fluctuations within one year (sudden response). We observed significant variation among provenances at all time-scales, but with varying degree among species. However, variation in short-term response (drought years) was remarkably unstable across all species, when the seasonal variation of drought occurrence was considered. Silver-fir and Douglas-fir showed significant associations between seed-source climate and growth response as well as trade-offs between early- and latewood growth reaction which strongly suggests that growth responses are adaptive. Intra-specific variation in conifers in response to drought will probably be sufficient to mitigate climate change consequences on forest growth, but growth-environment interactions as well as dependencies between temporal scales could create major pitfalls for adaptive forest management in the future.

Pre-fire drought and competition mediate post-fire conifer mortality in western U.S. National Parks.

Tree mortality is an important outcome of many forest fires. Extensive tree injuries from fire may lead directly to mortality, but environmental and biological stressors may also contribute to tree death. However, there is little evidence showing how the combined effects of two common stressors, drought and competition, influence post-fire mortality. Geographically broad observations of three common western coniferous trees subjected to prescribed fire showed the likelihood of post-fire mortality was related to intermediate-term (10 yr) pre-fire average radial growth, an important component of tree vigor. Path analysis showed that indices of competition and drought stress prior to fire can be described in terms of joint effects on growth, indirectly affecting post-fire mortality. Our results suggest that the conditions that govern the relationship between growth and mortality in unburned stands may also apply to post-fire environments. Thus, biotic and abiotic changes that affect growth negatively (e.g., drought stress) or positively (e.g., growth releases following thinning treatments) prior to fire may influence expressed fire severity, independent of fire intensity (e.g., heat flux, residence time). These relationships suggest that tree mortality may increase under stressful climatic or stand conditions even if fire behavior remains constant.

Two-stage patterning dynamics in conifer cotyledon whorl morphogenesis.

Background and Aims: Conifer embryos, unlike those of monocots or dicots, have variable numbers of cotyledons, even within the same species. Cotyledons form in a single whorl on a dome-shaped embryo. The closely spaced cotyledons are not found outside this ring, indicating a radial control on where they can form. Polar transport of the hormone auxin affects outgrowth of distinct cotyledons, but not the radial aspect of the whorl or the within-whorl spacing between cotyledons. A quantitative model of plant growth regulator patterning is needed to understand the dynamics of this complex morphogenetic process. Methods: A two-stage reaction-diffusion model is developed for the spatial patterning of growth regulators on the embryo surface, with a radial pattern (P1) constraining the shorter-wavelength cotyledon pattern (P2) to a whorl. These patterns drive three-dimensional (3-D) morphogenesis by catalysing local surface growth. Key Results: Growth driven by P2 generates single whorls across the experimentally observed range of two to 11 cotyledons, as well as the circularly symmetric response to auxin transport interference. These computations are the first corroboration of earlier theoretical proposals for hierarchical control of whorl formation. The model generates the linear relationship between cotyledon number and embryo diameter observed experimentally. This accounts for normal integer cotyledon number selection, as well as the less common cotyledon fusings and splittings observed experimentally. Flattening of the embryo during development may affect the upward outgrowth angle of the cotyledons. Conclusions: Cotyledon morphogenesis is more complex geometrically in conifers than in angiosperms, involving 2-D patterning which deforms a surface in three dimensions. This work develops a quantitative framework for understanding the growth and patterning dynamics involved in conifer cotyledon development, and applies more generally to the morphogenesis of whorls with many primordia.

Insect herbivory (Choristoneura fumiferana, Tortricidea) underlies tree population structure (Picea glauca, Pinaceae).

Variation in insect herbivory can lead to population structure in plant hosts as indicated by defence traits. In annual herbaceous, defence traits may vary between geographic areas but evidence of such patterns is lacking for long-lived species. This may result from the variety of selection pressures from herbivores, long distance gene flow, genome properties, and lack of research. We investigated the antagonistic interaction between white spruce (Picea glauca) and spruce budworm (SBW, Choristoneura fumiferana) the most devastating forest insect of eastern North America in common garden experiments. White spruces that are able to resist SBW attack were reported to accumulate the acetophenones piceol and pungenol constitutively in their foliage. We show that levels of these acetophenones and transcripts of the gene responsible for their release is highly heritable and that their accumulation is synchronized with the most devastating stage of SBW. Piceol and pungenol concentrations negatively correlate with rate of development in female SBW and follow a non-random geographic variation pattern that is partially explained by historical damage from SBW and temperature. Our results show that accumulation of acetophenones is an efficient resistance mechanism against SBW in white spruce and that insects can affect population structure of a long-lived plant.

Phenological research of climate changes in the north part of Lithuania by the phenological garden of Siauliai University.

Siauliai University Botanical Garden is a member of the International Phenological Garden network since 2005. It is the only one botanical garden in the East Europe that participated in the programme. In 2015, 18 species were observed. For research, data of 14 plants was used. The aim of this study is to estimate the responsiveness of the species of plants of the phenological garden to annual and monthly precipitation and temperature of the air. The main variables in this investigation were growing season length and the beginning of the growing season. In the period 2006-2015, the lowest annual air temperature was in 2010 (6.0 °C), and the highest was in 2015 (8.9 °C). The lowest precipitation was in 2015 (37.3 mm), and the highest was in 2012 (63.5 mm). The leanest regression among growing length, average annual precipitation, and air temperature showed that statistically significant correlation between growing length and average annual air temperature was found for nine plants, between growing length and precipitation was found for three plants, and between growing length and both factors was found for one plant, Salix smithiana, only. Due to the short evaluating period (2007-2015), consistent regression of the length of the growing season could not be found. The growing length of Betula pubescens sequentially increased. The average growing season of 14 plants starts on April 27 (±3), but for Corylus avellana, it is on April 26 (±3). Longevity of the growing season was the most related with precipitation for C. avellana in summer, autumn, and winter and with air temperature, Ribes alpinum and Salix acutifolia in summer and in autumn.

Climate-Driven Synchronized Growth of Alpine Trees in the Southeast Tibetan Plateau.

Knowledge about the spatiotemporal tree growth variability and its associations with climate provides key insights into forest dynamics under future scenarios of climate change. We synthesized 17 tree-ring width chronologies from four tree species at the high-elevation sites in the southeast Tibetan Plateau (SETP) to study the regional tree growth variability and climate-growth relationships. Despite of diverse habitats and different physiological characteristics of these species, these tree-ring chronologies shared a significant common variance in SETP. An unprecedented increase in the shared variance is found along the latter half of the 20th century, coinciding with the enhancement of the frequency of extreme rings among chronologies. It is found that minimum winter temperature tends to be the dominant climate for trees in this region. The site-specific responses in cold (1965-1980) and warm (1990-2005) intervals by means of Fuzzy Cmeans (FCM) clustering reveal that the remarkable enhancement of growth synchrony among trees mainly occur in warm conditions. This is different from previous findings indicating that increased consistence among temperature sensitive tree rings in cold periods. This may be related to the reduced temperature sensitivity of regional tree growth as winter minimum temperature is lower than a certain threshold, which is in agreement with the "principle of ecological amplitude". In addition, it is worth noting that precipitation in June have started to restrain the tree growth since the beginning of the 1980s, which is possibly an important contributor for synchronized growth among trees in SETP.

Ectomycorrhizas in vitro between Tricholoma matsutake, a basidiomycete that associates with Pinaceae, and Betula platyphylla var. japonica, an early-successional birch species, in cool-temperate forests.

Tricholoma matsutake is an ectomycorrhizal basidiomycete that associates with Pinaceae in the Northern Hemisphere and produces prized "matsutake" mushrooms. We questioned whether the symbiont could associate with a birch that is an early-successional species in boreal, cool-temperate, or subalpine forests. In the present study, we demonstrated that T. matsutake can form typical ectomycorrhizas with Betula platyphylla var. japonica; the associations included a Hartig net and a thin but distinct fungal sheath, as well as the rhizospheric mycelial aggregate "shiro" that is required for fruiting in nature. The in vitro shiro also emitted a characteristic aroma. This is the first report of an ectomycorrhizal formation between T. matsutake and a deciduous broad-leaved tree in the boreal or cool-temperate zones that T. matsutake naturally inhabits.

Sapling leaf trait responses to light, tree height and soil nutrients for three conifer species of contrasting shade tolerance.

We developed models to describe the responses of four commonly examined leaf traits (mass per area, weight, area and nitrogen (N) concentration) to gradients of light, soil nutrients and tree height in three conifer species of contrasting shade tolerance. Our observational dataset from the sub-boreal spruce forests of British Columbia included subalpine fir (Abies lasioscarpa [Hook.] Nutt; high shade tolerance), interior spruce (Picea glauca × Picea engelmannii [Moench] Voss; intermediate shade tolerance) and lodgepole pine (Pinus contorta Dougl. ex Loud. var. latifolia; low shade tolerance) saplings from 0.18 to 4.87 m tall, in 8-98% of total incident light, from field sites with <17.6 kg ha(-1) to >46.8 kg ha(-1) total dissolved N. Leaf weights and areas showed strong positive responses to light and height, but little or no response to soil nutrients. Parameter estimates indicated that the shape of leaf weight and area responses to light corresponded with shade tolerance ranking for the three species; pine had the most linear response whereas spruce and fir had asymptotic responses. Leaf N concentration responded positively to soil nutrients, negatively to light and idiosyncratically to height. The negative effect of light was only apparent on sites of high soil nutrient availability, and parameter estimates for the shape of the negative response also corresponded to shade tolerance ranking (apine = -0.79, aspruce = -0.15, afir = -0.07). Of the traits we measured, leaf mass per area showed the least response to light, soil nutrient and height gradients. Although it is a common practice in comparisons across many species, characterizing these conifers by mean values of their leaf traits would miss important intraspecific variation across environmental and size gradients. In these forests, parameter estimates representing the intraspecific variability of leaf trait responses can be used to understand relative shade tolerances.

[Physical factors influencing the floristic relationships of pinyon pine (Pinaceae) from San Luis Potosi, Mexico]. / Factores físicos que influyen en las relaciones florísticas de los piñonares (Pinaceae) de San Luis Potosí, México.

In plant communities, the species distribution patterns and their relationships with environmental factors are of central importance in ecology. In San Luis Potosí of Mexico, woodlands of Pinus cembroides and P. johannis are sympatric, but P. cembroides tends to be located in South and Southwest slopes, in more disturbed sites; unlike, P. johannis, is mostly distributed in mesic areas, in North and Northeast slopes. The aim of this study was to analyze the influence of some physical factors on the floristic similarity of pinyon pine P. cembroides and P. johannis. The study area was located in the Sierra San Miguelito, San Luis Potosí, Mexico. We selected 40 sampling units spread over an area of 50km2. In each unit, we laid out two 20m long lines perpendicular to each other, in which we recorded cover data of the plant species intercepted. We developed two data matrices, the first one including cover values of 91 species, and the second one, considering seven topographical, climatic, and solar radiation variables. We applied cluster analysis and ordination to explore the influence of environmental variables on the floristic differentiation of pinyon pine woodlands. Clustering showed six groups, the first three characterized by P. cembroides. The ordination showed that variance represented by the first three axes was 65.9%. Axis 1 was positively correlated with altitude and negatively with mean annual temperature; axes two and three, showed low correlation with the variables tested. P. cembroides woodlands and accompanying flora tend to be located in lower altitude, higher mean annual temperature, and mainly in South-Southwestern slopes. In contrast, stands of P. johannis, mixed stands of P. johannis-P. cembroides, and Quercus potosina, were usually founded in greater altitudes, mean annual temperature slightly lower, and North-Northeastern exposure. The sites of these monospecific and mixed woodlands with associated species, indicators of environmental variables, generates the pre-diagnosis of the situation of the communities and their condition, so that future actions can be planned in agreement with the natural balance of the ecosystem.

Factores físicos que influyen en las relaciones florísticas de los piñonares (Pinaceae) de San Luis Potosí, México / Physical factors influencing the floristic relationships of pinyon pine (Pinaceae) from San Luis Potosi, Mexico

In plant communities, the species distribution patterns and their relationships with environmental factors are of central importance in ecology. In San Luis Potosi of Mexico, woodlands of Pinus cembroides and P. johannis are sympatric, but P. cembroides tends to be located in South and Southwest slopes, in more disturbed sites; unlike, P. johannis, is mostly distributed in mesic areas, in North and Northeast slopes. The aim of this study was to analyze the influence of some physical factors on the floristic similarity of pinyon pine P. cembroides and P. johannis. The study area was located in the Sierra San Miguelito, San Luis Potosi, Mexico. We selected 40 sampling units spread over an area of 50km². In each unit, we laid out two 20m long lines perpendicular to each other, in which we recorded cover data of the plant species intercepted. We developed two data matrices, the first one including cover values of 91 species, and the second one, considering seven topographical, climatic, and solar radiation variables. We applied cluster analysis and ordination to explore the influence of environmental variables on the floristic differentiation of pinyon pine woodlands. Clustering showed six groups, the first three characterized by P. cembroides. The ordination showed that variance represented by the first three axes was 65.9%. Axis 1 was positively correlated with altitude and negatively with mean annual temperature; axes two and three, showed low correlation with the variables tested. P. cembroides woodlands and accompanying flora tend to be located in lower altitude, higher mean annual temperature, and mainly in South-Southwestern slopes. In contrast, stands of P. johannis, mixed stands of P. johannis-P. cembroides, and Quercus potosina, were usually founded in greater altitudes, mean annual temperature slightly lower, and North-Northeastern exposure. The sites of these monospecific and mixed woodlands with associated species, indicators of environmental variables, generates the pre-diagnosis of the situation of the communities and their condition, so that future actions can be planned in agreement with the natural balance of the ecosystem.

El conocimiento de los patrones de distribución de especies en las comunidades vegetales y su relación con factores del medio es un objetivo fundamental en ecología. Las comunidades de Pinus cembroides y P. johannis coexisten en diversas regiones de México, pero P. cembroides tiende a distribuirse en laderas de exposiciones sur y suroeste en sitios deteriorados, a diferencia de P. johannis localizados en áreas mésicas en laderas con exposición norte y noreste. El objetivo de este trabajo fue analizar la influencia de algunos factores físicos en la semejanza florística de los piñonares de P. cembroides y P. johannis. El área de estudio se ubica en la Sierra San Miguelito, San Luis Potosí, México. Se seleccionaron 40 unidades de muestreo, distribuidas en una superficie de 50km²; en cada una se trazaron dos líneas de 20m perpendiculares entre sí, donde se registró la cobertura de los individuos interceptados. Se elaboraron dos tipos de matrices de datos: la primera con los valores de cobertura de 91 especies y la segunda, con siete variables topográficas, climáticas y de incidencia solar. Se aplicaron análisis de agrupamiento y de ordenación para explorar la influencia de variables ambientales en la diferenciación florística de los piñonares. El agrupamiento diferenció seis grupos; los tres primeros caracterizados por bosques de P. cembroides. La ordenación mostró que la varianza representada por los tres primeros ejes fue de 65.9%. El eje uno se correlacionó positivamente con la altitud y negativamente con la temperatura media anual y la orientación; los ejes dos y tres mostraron bajos valores de correlación con las variables probadas. Se observó que los piñonares de P. cembroides y flora acompañante, tienden a localizarse en sitios de menor altitud, con mayor temperatura media anual y en laderas con orientación al sur y suroeste. En contraste, los sitios de P. johannis, los de P. johannis-P. cembroides y de éstos con Quercus potosina, por lo regular están ubicados en condiciones de mayor altitud, temperatura media anual baja y con orientación al norte-noreste. La localización de estos bosques monoespecíficos y mixtos junto con las especies asociadas, indicadoras de variables ambientales, genera el pre-diagnóstico de la situación de las comunidades y su condición, para que en el futuro se planifiquen acciones acordes con el balance natural ecosistémico.

Trade-offs between constitutive and induced defences drive geographical and climatic clines in pine chemical defences.

There is increasing evidence that geographic and climatic clines drive the patterns of plant defence allocation and defensive strategies. We quantified early growth rate and both constitutive and inducible chemical defences of 18 Pinaceae species in a common greenhouse environment and assessed their defensive allocation with respect to each species' range across climatic gradients spanning 31° latitude and 2300 m elevation. Constitutive defences traded-off with induced defences, and these defensive strategies were associated with growth rate such that slow-growing species invested more in constitutive defence, whereas fast-growing species invested more in inducible defence. The position of each pine species along this trade-off axis was in turn associated with geography; moving poleward and to higher elevations, growth rate and inducible defences decreased, while constitutive defence increased. These geographic patterns in plant defence were most strongly associated with variation in temperature. Climatic and geographical clines thus act as drivers of defence profiles by mediating the constraints imposed by trade-offs, and this dynamic underlays global patterns of defence allocation.

Drought response of five conifer species under contrasting water availability suggests high vulnerability of Norway spruce and European larch.

The ability of tree species to cope with anticipated decrease in water availability is still poorly understood. We evaluated the potential of Norway spruce, Scots pine, European larch, black pine, and Douglas-fir to withstand drought in a drier future climate by analyzing their past growth and physiological responses at a xeric and a mesic site in Central Europe using dendroecological methods. Earlywood, latewood, and total ring width, as well as the δ(13) C and δ(18) O in early- and latewood were measured and statistically related to a multiscalar soil water deficit index from 1961 to 2009. At the xeric site, δ(13) C values of all species were strongly linked to water deficits that lasted longer than 11 months, indicating a long-term cumulative effect on the carbon pool. Trees at the xeric site were particularly sensitive to soil water recharge in the preceding autumn and early spring. The native species European larch and Norway spruce, growing close to their dry distribution limit at the xeric site, were found to be the most vulnerable species to soil water deficits. At the mesic site, summer water availability was critical for all species, whereas water availability prior to the growing season was less important. Trees at the mesic were more vulnerable to water deficits of shorter duration than the xeric site. We conclude that if summers become drier, trees growing on mesic sites will undergo significant growth reductions, whereas at their dry distribution limit in the Alps, tree growth of the highly sensitive spruce and larch may collapse, likely inducing dieback and compromising the provision of ecosystem services. However, the magnitude of these changes will be mediated strongly by soil water recharge in winter and thus water availability at the beginning of the growing season.

Rapid warming accelerates tree growth decline in semi-arid forests of Inner Asia.

Forests around the world are subject to risk of high rates of tree growth decline and increased tree mortality from combinations of climate warming and drought, notably in semi-arid settings. Here, we assess how climate warming has affected tree growth in one of the world's most extensive zones of semi-arid forests, in Inner Asia, a region where lack of data limits our understanding of how climate change may impact forests. We show that pervasive tree growth declines since 1994 in Inner Asia have been confined to semi-arid forests, where growing season water stress has been rising due to warming-induced increases in atmospheric moisture demand. A causal link between increasing drought and declining growth at semi-arid sites is corroborated by correlation analyses comparing annual climate data to records of tree-ring widths. These ring-width records tend to be substantially more sensitive to drought variability at semi-arid sites than at semi-humid sites. Fire occurrence and insect/pathogen attacks have increased in tandem with the most recent (2007-2009) documented episode of tree mortality. If warming in Inner Asia continues, further increases in forest stress and tree mortality could be expected, potentially driving the eventual regional loss of current semi-arid forests.

Disparity in elevational shifts of European trees in response to recent climate warming.

Predicting climate-driven changes in plant distribution is crucial for biodiversity conservation and management under recent climate change. Climate warming is expected to induce movement of species upslope and towards higher latitudes. However, the mechanisms and physiological processes behind the altitudinal and latitudinal distribution range of a tree species are complex and depend on each tree species features and vary over ontogenetic stages. We investigated the altitudinal distribution differences between juvenile and adult individuals of seven major European tree species along elevational transects covering a wide latitudinal range from southern Spain (37°N) to northern Sweden (67°N). By comparing juvenile and adult distributions (shifts on the optimum position and the range limits) we assessed the response of species to present climate conditions in relation to previous conditions that prevailed when adults were established. Mean temperature increased by 0.86 °C on average at our sites during the last decade compared with previous 30-year period. Only one of the species studied, Abies alba, matched the expected predictions under the observed warming, with a maximum abundance of juveniles at higher altitudes than adults. Three species, Fagus sylvatica, Picea abies and Pinus sylvestris, showed an opposite pattern while for other three species, such as Quercus ilex, Acer pseudoplatanus and Q. petraea, we were no able to detect changes in distribution. These findings are in contrast with theoretical predictions and show that tree responses to climate change are complex and are obscured not only by other environmental factors but also by internal processes related to ontogeny and demography.

Climate determines upper, but not lower, altitudinal range limits of Pacific Northwest conifers.

Does climate determine species' ranges? Rapid rates of anthropogenic warming make this classic ecological question especially relevant. We ask whether climate controls range limits by quantifying relationships between climatic variables (precipitation, temperature) and tree growth across the altitudinal ranges of six Pacific Northwestern conifers on Mt. Rainier, Washington, USA. Results for three species (Abies amabilis, Callitropsis nootkatensis, Tsuga mertensiana) whose upper limits occur at treeline (> 1600 m) imply climatic controls on upper range limits, with low growth in cold and high snowpack years. Annual growth was synchronized among individuals at upper limits for these high-elevation species, further suggesting that stand-level effects such as climate constrain growth more strongly than local processes. By contrast, at lower limits climatic effects on growth were weak for these high-elevation species. Growth-climate relationships for three low-elevation species (Pseudotsuga menziesii, Thuja plicata, Tsuga heterophylla) were not consistent with expectations of climatic controls on upper limits, which are located within closed-canopy forest (< 1200 m). Annual growth of these species was poorly synchronized among individuals. Our results suggest that climate controls altitudinal range limits at treeline, while local drivers (perhaps biotic interactions) influence growth in closed-canopy forests. Climate-change-induced range shifts in closed-canopy forests will therefore be difficult to predict accurately.

Yields and constituents of essential oil from cones of Pinaceae spp. natively grown in Turkey.

In this study, the yields and composition of essential oils obtained from the cones of Pinaceae family species natively grown in Turkey were investigated. Essential oils were obtained by hydrodistillation. Oil yields were 0.13-0.48 mL/100 g in pine cones, 0.42-0.59 mL/100g in fir, 0.36 mL/100g in spruce and 0.37 mL/100g in cedar. While alpha-pinene (47.1-14.8%) was the main constituent of P. slyvestris, P. nigra and P. halepensis, limonene (62.8%) in P. pinea and beta-pinene (39.6%) in P. brutia were found in higher amounts. Like in P. pinea, limonene was the main compound in Cedrus libani (22.7%). In fir species the major compounds were alpha-pinene (70.6-53.0%) and beta-pinene (10.9-8.2%). Contrary to other species beta-pinene (32.7%) was found as a major compound in Picea orientalis.

A novel insight into the regulation of light-independent chlorophyll biosynthesis in Larix decidua and Picea abies seedlings.

Light-independent chlorophyll (Chl) biosynthesis is a prerequisite for the assembly of photosynthetic pigment-protein complexes in the dark. Dark-grown Larix decidua Mill. seedlings synthesize Chl only in the early developmental stages and their Chl level rapidly declines during the subsequent development. Our analysis of the key regulatory steps in Chl biosynthesis revealed that etiolation of initially green dark-grown larch cotyledons is connected with decreasing content of glutamyl-tRNA reductase and reduced 5-aminolevulinic acid synthesizing capacity. The level of the Chl precursor protochlorophyllide also declined in the developing larch cotyledons. Although the genes chlL, chlN and chlB encoding subunits of the light-independent protochlorophyllide oxidoreductase were constitutively expressed in the larch seedlings, the accumulation of the ChlB subunit was developmentally regulated and ChlB content decreased in the fully developed cotyledons. The efficiency of chlB RNA-editing was also reduced in the mature dark-grown larch seedlings. In contrast to larch, dark-grown seedlings of Picea abies (L.) Karst. accumulate Chl throughout their whole development and show a different control of ChlB expression. Analysis of the plastid ultrastructure, photosynthetic proteins by Western blotting and photosynthetic parameters by gas exchange and Chl fluorescence measurements provide additional experimental proofs for differences between dark and light Chl biosynthesis in spruce and larch seedlings.

Anchorage of mature conifers: resistive turning moment, root-soil plate geometry and root growth orientation.

Eighty-four mature Norway spruce (Picea abies L. Karst), silver fir (Abies alba Mill) and Scots pine (Pinus sylvestris L.) trees were winched over to determine the maximum resistive turning moment (M(a)) of the root-soil system, the root-soil plate geometry, the azimuthal orientation of root growth, and the occurrence of root rot. The calculation of M(a), based on digital image tracking of stem deflection, accounted not only for the force application and its changing geometry, but also for the weight of the overhanging tree, representing up to 42% of M(a). Root rot reduced M(a) significantly and was detected in 25% of the Norway spruce and 5% of the silver fir trees. Excluding trees with root rot, differences in M(a) between species were small and insignificant. About 75% of the variance in M(a) could be explained by one of the four variables--tree mass, stem mass, stem diameter at breast height squared times tree height, and stem diameter at breast height squared. Among the seven allometric variables assessed above ground, stem diameter at breast height best described the root-soil plate dimensions, but the correlations were weak and the differences between species were insignificant. The shape of the root-soil plate was well described by a depth-dependent taper model with an elliptical cross section. Roots displayed a preferred azimuthal orientation of growth in the axis of prevailing winds, and the direction of frequent weak winds matched the orientation of growth better than that of rare strong winds. The lack of difference in anchorage parameters among species probably reflects the similar belowground growth conditions of the mature trees.