Which factors most influence heartwood distribution and radial growth in plantation teak?

? To investigate the effect of climate on radial growth in young plantation grown teak (Tectona grandis L.), growth ring width was measured in 105 trees and correlated to precipitation and temperature data.

? The social status of trees within the stand was also determined and cross-sectional area (CSA) for the trunk correlated to the proportion of heartwood (HW) within the tree. HW develops asymmetrically in leaning stems of some conifer species, but it is not known if this phenomenon also occurs in broadleaf species. Therefore, we measured HW proportion in leaning and straight stems, along with the number of growth rings in the HW.

? Annual ring width depended strongly on mean monthly temperature during the rainy season and the most significant relationships were found corresponding to the months of June and July. With regard to the weaker relationship between precipitation and radial growth, correlations were highest during the period of bud-break at the beginning of the rainy season.

? The very high stand density affected radial growth, particularly in suppressed trees, which responded little to thinning operations. HW formation was greatest in dominant trees, and was highly regressed with stem CSA.

? Therefore, rapid growth of young stands should be encouraged by reducing stand density. Asymmetric HW formation occurred in both leaning and straight trees, and was significantly greater along the upper sides of leaning stems. It is probable that this eccentric HW formation is linked to mechanical loading on the tree.

     

Large-scale structural variation and long-term growth dynamics of Juniperus thurifera trees in a managed woodland in Soria, central Spain

? Juniper woodlands have a great conservational and productive importance in Spain. However, basic traits of their structural variation and growth dynamics are unknown.

? To characterize the structural variation and assess both height and radial growth patterns, we sampled and measured stem cross-sections every 0.5 m in height from 107 Spanish juniper (Juniperus thurifera L.) trees, which were selected based on a 350 m regular grid over a whole woodland of 3 300 ha in area. We used dendroecological techniques and spatial analysis to study tree structure and growth.

? Structural traits such as bole diameter and height were mainly determined by tree life-related parameters such as tree age and growth rate variation, while abiotic factors such as altitude played a minor role in determining structural variation. Over the last 300 years, juniper establishment has been continuous in time but discontinuous in space. Large-scale spatial heterogeneity of tree establishment, and the presence of an early growth suppression and abrupt growth changes in junipers > 100 years old are consequences of an intense management that almost ceased in the late 19th century.

? Abrupt growth releases and suppressions were synchronic among different age classes, and coincided with documentary records on livestock decrease and key historical changes in land use. Growth patterns suggest that juniper is a brow sing-tolerant species that is able to survive large periods of intense browsing pressure.

     

Influence of annual weather on growth of pedunculate oak in southern Sweden

? A network of oak (Quercus robur L.) chronologies containing 49 sites and 635 single trees was analysed to identify weather variables affecting annual tree-ring increment dynamics in southern Sweden during 1860–2000.

? We analysed (1) the growth response of oak to non-extreme weather, and (2) the temporal and spatial patterns of regional growth anomalies (pointer years) and associated climatic extremes resolved on a monthly scale.

? Growth was controlled by precipitation in the current (June–July) and the previous growing season (August) in 48% and 22% of all sites, respectively. Temperature during July of the current year and August of the previous year was negatively correlated with growth in 29% and 43% of the sites, respectively. Growth was positively correlated with temperature in October of the previous season in 72% of the sites. The most extensive growth anomaly occurred in 1965 and was probably caused by intrusion of cold Arctic air masses into the region at the end of March that year.

? During climatically non-extreme years, oak growth is driven mostly by the dynamics of summer precipitation. Many of the negative growth anomalies, however, were associated with temperature extremes. Southern Swedish oak pointer years tend not to coincide with the pan-European oak pointer years.

     

Effects of climate variables on intra-annual stem radial increment in Pinus cembra (L.) along the alpine treeline ecotone

? Within the alpine treeline ecotone tree growth is increasingly restricted by extreme climate conditions. Although intra-annual stem growth recorded by dendrometers can be linked to climate, stem diameter increments in slow-growing subalpine trees are masked by changes in tree water status.

? We tested the hypothesis that intra-annual radial stem growth in Pinus cembra is influenced by different climate variables along the treeline ecotone in the Austrian Alps. Dendrometer traces were compared with dynamics of xylem cell development to date onset of cambial activity and radial stem growth in spring.

? Daily fluctuations in stem radius reflected changes in tree water status throughout the treeline ecotone. Extracted daily radial increments were significantly correlated with air temperature at the timberline and treeline only, where budburst, cambial activity and enlargement of first tracheids also occurred quite similarly. A close relationship was detected between radial increment and number of enlarging tracheids throughout the treeline ecotone.

? We conclude that (i) the relationship between climate and radial stem growth within the treeline ecotone is dependent on a close coupling to atmospheric climate conditions and (ii) initiation of cambial activity and radial growth in spring can be distinguished from stem re-hydration by histological analysis.

     

Growth response of Pinus pinaster Ait. to climatic variables in central Spanish forests

? The effect of climatic variables (temperature and precipitation) on radial growth of the Mediterranean Maritime pine (Pinus pinaster Ait.) was studied using dendrochronological techniques in the Iberian Peninsula.

? Ten tree-ring width chronologies, along the central distribution area of the species, were built. Chronology variability was analysed using Principal Component Analysis (PCA) for the period 1952–2005.

? The first principal component (PC1) explained 56% of tree-growth variability. Tree-growth association with climate was analysed at regional and local scales using correlation coefficient and bootstrapped response functions.

? Radial growth at both scales was positively correlated with rainfall during and prior to the growing season at all sites, and with summer rainfall before the growing season at five sites. Mean temperature effect changed according to the sampling site, from non-significant at the highest sites to significant (positive relationship in winter) at the lowest sites. Growth season temperature also had a negative effect.

? The Kalman filter was applied to estimate changing association between growth and climate over-time. Results suggested a change in association, initiated in the 80s, from non-significant to significant (*p < 0.05) at six of the sampling sites.

? Pinus pinaster is an accurate species for analysing tree-growth association with climate and for studying plant behaviour under global change conditions.

     

Black pine (Pinus nigra Arn.) growth divergence along a latitudinal gradient in Western Mediterranean mountains

? Most studies of tree-growth and climate report positive responses to global warming in high latitudes and negative responses at lower ones.

? We analyzed tree-ring width of Pinus nigra Arn. along a 500 km latitudinal transect in the Iberian Peninsula to study the temporal trend and climate forcing in tree radial growth during the last century.

? Tree growth was enhanced by cool summers and moist cold seasons. Increased moisture stress has decreased tree growth rates. However, we present evidence of growth increases in some trees in all sampled populations after 1980’s. Climate change negatively (positively) affected between 72% (5%) of trees in the southern populations and 40% (25%) in the north Trees with positive growth trends were favored by winter temperatures and their abundance was inversely correlated with forest productivity.

? Our findings add evidences of tree growth divergence in the Mediterranean basin and show the gradual transition between forests where positive (temperate and boreal) and negative (Mediterranean) growth trends dominate.

     

Has global change induced divergent trends in radial growth of Pinus sylvestris and Pinus halepensis at their bioclimatic limit? The example of the Sainte-Baume forest (south-east France)

? It is agreed that climate (precipitation and temperature) influences the distribution of plant species. Near the margins of a species’ natural range, climate becomes limiting to physiological processes. There, climate change may be expected to have a significant impact on tree growth and the species’ ranges may be altered.

? In order to assess what influence climate change could exert on the distribution of pine species at their margin, radial growth trends in ring-width chronologies over the last century were analysed. In the French Mediterranean area where climate change is characterized by increased temperature, forest plots were selected along an altitudinal transect on the north-facing slope of the Sainte-Baume mountain (Bouches-du-Rhône, France) where the ranges of Pinus sylvestris and Pinus halepensis overlap.

? Two growth patterns were identified. For P. halepensis, radial growth has increased at all altitudes indicating that climate change has improved growth conditions of this species near the margin of its ecological range. For P. sylvestris, radial growth has increased only at low altitudes and even decreased at high altitudes.

? It must be deduced that the growth changes observed cannot be generalised either at the species level or at the geographical level and must be interpreted with great caution.

     

Climate-growth variability in Quercus ilex L. west Iberian open woodlands of different stand density

? We present the longest tree-ring chronology (141 y) of Quercus ilex L. (holm oak), and discuss the species climate-growth relationships and the influence of stand density on tree sensitivity to climate.

? Similarly to Quercus suber L., the most influential climatic variables upon holm oak growth were late spring and early summer precipitation, which enhanced growth, and high temperatures in the previous August and current July, which negatively affected growth.

? High density stands responded to similar climatic factors as low density stands, but their response was generally weaker. Holm oak sensitivity to climate has increased in recent decades, which might be related to increasing temperatures in the region. Sensitivity was higher in low density stands. Additionally, the effect of summer stress on growth seems to have increased during the same period, similarly to other species in the Iberian Peninsula, suggesting that trees are more vulnerable to climatic changes.

? Stand density could buffer the response to climate by smoothing climatic extremes. Nevertheless, the effect of competition might reverse this positive effect at the individual tree level. Precautions should be taken before providing management guidelines regarding the effect of climate change and stand density on holm oak.

     

High resolution temporal variation in wood properties in irrigated and nonirrigated Eucalyptus globulus

? Environmental determinants of wood properties variation were examined in Eucalyptus globulus, a globally important hardwood plantation species, in southern Tasmania, Australia.

? Radial variation in wood properties, measured with the SilviScan system, were re-scaled from distance to time abscissa using stem radial growth data measured with dendrometers. With this re-scaled data it was possible to evaluate how water availability and temperature affected wood density, microfibril angle (MFA) and fibre and vessel transverse dimensions in irrigated and non-irrigated trees.

? Wood density, fibre radial diameter and MFA were sensitive to water availability. Wood density increased and fibre radial diameter decreased in response to reduced water availability. When high water availability was maintained, wood density was negatively correlated with temperature. Together, temperature and soil matric potential explained about 60% of temporal variation in wood density variation. In contrast MFA was not related to temperature but decreased with increasing water stress. Slower growing trees also had lower MFA than faster growing trees. Slower growing trees had a larger number of vessels per unit area of wood than faster growing trees within this even aged stand. However, vessel radius to the 4th power was significantly higher in faster growing trees than in slower growing trees.

? Overall, E. globulus wood properties were sensitive to temporal changes in environmental conditions (particularly water availability) and associated growth rates. The data provided support for the hypothesis that growth rates are hydraulically mediated.

     

Effects of organic residue management and legume cover on growth of pine seedlings, nutrient leaching and soil properties

? The short-term effect of organic residue management on the growth and nutrition of Pinus pinaster Ait. seedlings, and on nutrient leaching and chemical properties of an acid soil was assessed through a lysimeter experiment. Treatments included absence, placement on the soil surface, and incorporation into the soil (with and without legume cover cropping) of organic residues (forest floor litter or forest floor litter plus harvest residues).

? Residues placed on the soil surface enhanced seedling growth. Organic residues reduced nutrient losses (NO ₃ ^? , Ca and Mg) and resulted in nutrient accumulation in the soil. Harvest residues positively affected K seedling nutrition status and enhanced K soil accumulation.

? Legume cover cropping reduced soil nutrient losses (N, Ca, Mg and K) during the early stage of seedling growth; it also improved seedling nutrition status (N and P), but without any effect on growth.

? Harvest residues plus forest floor litter placed on the soil surface was the most appropriate management to both reduce nutrient losses through leaching and increase height of seedlings at the end of the experimental period (two years).

     

Is forest mushroom productivity driven by tree growth? Results from a thinning experiment

? Most of the edible forest mushrooms are mycorrhizal and depend on carbohydrates produced by the associated trees. Fruiting patterns of these fungi are not yet fully understood since climatic factors alone do not completely explain mushroom occurrence.

? The objective of this study was to retrospectively find out if changing tree growth following an increment thinning has influenced the diversity patterns and productivity of associated forest mushrooms in the fungus reserve La Chanéaz, Switzerland.

? The results reveal a clear temporal relationship between the thinning, the growth reaction of trees and the reaction of the fungal community, especially for the ectomycorrhizal species. The tree-ring width of the formerly suppressed beech trees and the fruit body number increased after thinning, leading to a significantly positive correlation between fruit body numbers and tree-ring width.

? Fruit body production was influenced by previous annual tree growth, the best accordance was found between fruit body production and the tree-ring width two years previously.

? The results support the hypothesis that ectomycorrhizal fruit body production must be linked with the growth of the associated host trees. Moreover, the findings indicate the importance of including mycorrhizal fungi as important players when discussing a tree as a carbon source or sink.

     

Cold hardiness and transplant response of Juglans nigra seedlings subjected to alternative storage regimes

? Effects of overwinter storage regimes on seedling cold hardiness and physiological vigor are relatively unexplored, particularly for temperate deciduous forest tree species.

? We evaluated influence of storage duration (0, 66, 119, or 175 d) on electrolyte leakage of stem and root collar tissues following exposure to a series of freeze-test temperatures in black walnut (Juglans nigra L.) seedlings sampled from cold (3 °C) or freezer (?2 °C) storage. Seedlings were subsequently transplanted into a controlled growth chamber environment for two months.

? Regardless of storage temperature, mean LT₅₀ was lowest for seedlings stored for 66 d (≤ ?34 °C) and increased dramatically after 119 d (≥ ?13 °C).

? Root collar tissue had lower LT₅₀ than stem tissue after 119 d for cold-stored seedlings, reflecting importance of evaluative tissue type. Days to bud break shortened with increasing storage duration up to 119 d and stabilized thereafter for both storage regimes. Root growth potential was maximized after 119 d of storage, and subsequently declined for cold-stored seedlings. Height growth increased following storage, regardless of duration.

? To promote stress resistance and transplant growth response, we recommend that black walnut seedlings from this genetic source be outplanted after approximately 66–119 d of storage.

     

Photosynthetic temperature adaptation of Pinus cembra within the timberline ecotone of the Central Austrian Alps

? Temperature is suggested to determine the upper limit of tree life. Therefore, future climate warming may be of importance for tree distribution within the European Alps, where low temperatures limit carbon metabolism.

? We focused on the effects of air and soil temperature on net photosynthesis (P _n) of Pinus cembra an evergreen climax species of the timberline ecotone of the Central Austrian Alps. Light response and temperature response curves were estimated along an altitudinal gradient ranging from the forest limit up to the krummholz limit in both summer and fall.

? In general, P _n was significantly lower in fall as compared to summer. Nevertheless, independent from season mean P _n values tended to increase with elevation and were positively correlated with root zone temperatures. The specific leaf area by contrast declined with increasing elevation. Furthermore, the temperature optimum of net photosynthesis declined with increasing elevation and was positively correlated with the mean maximum air temperature of the 10 days prior the date of measurement.

? Thus, our findings appear to reflect a long-term adaptation of the photosynthetic apparatus of Pinus cembra to the general temperature conditions with respect to elevation combined with a short term acclimation to the prevailing temperature regime.

     

Effects of site on fibre, kraft pulp and handsheet properties of Eucalyptus globulus

? Eight-year old trees from two Eucalyptus globulus Labill. clones planted across three different sites in Tasmania, Australia, were sampled for wood and kraft pulp/handsheet properties.

? Site had a significant effect on all measured properties. Compared with the poor site (Parkham) the wood from the good site (West Ridgley) had on average 11 % lower wood density. The poor site had also greater microfibril angles, shorter fibres at lower pulp yields.

? The handsheets produced with pulp from the poor site resulted in comparatively higher bulkiness, lower burst, lower tear and tensile indices, lower zero span tensile strength, but higher opacity, higher light scattering and higher surface roughness. Significant height effects were found with all wood properties, and also with tear index, zero span tensile strength and opacity.

? Discriminant analysis showed that for 76 out of 100 handsheets the raw material source, i.e. growth site, could be predicted correctly using a set of handsheet properties with tear index and bulk index being most prominent.

? This is unique evidence that site conditions are strongly reflected in handsheet properties produced from Eucalyptus pulp.

     

Genetic variation for carbon isotope composition in Juglans regia L.: relationships with growth, phenology and climate of origin

? Among the traits of breeding interest for the common walnut tree Juglans regia L., characteristics such as timing of budbreak and leaf fall, water-use efficiency and growth performance are regarded as being of utmost relevance in Mediterranean conditions.

? The authors evaluated intraspecific variation in δ¹³C (carbon isotope composition, surrogate of intrinsic water-use efficiency, WUE_i) for 22 J. regia families grown in a progeny test under supplementary irrigation, and investigated whether such variation correlated with climatic indicators of native habitats. The genetic relationships between δ¹³C, growth and phenology were also assessed during two consecutive years.

? Overall, the most water-use-efficient families (i.e. with higher δ¹³C), which originated mainly from drought-prone provenance regions which have a high vapour pressure deficit and low rainfall, exhibited less height growth and smaller DBH. Using a stepwise regression procedure, δ¹³C was included as the main explanatory variable of genotypic variation in growth traits, together with growing season duration (for DBH in both years) and flushing (for height in 2007).

? It was concluded that WUE_i is largely unconnected to phenology effects in the explanation of growth performance for J. regia, therefore suggesting the opportunity of simultaneously selecting for low WUE_i and extended growing period to maximise productivity in non-water-limited environments.

     

Comparing needle and shoot phenology with xylem development on three conifer species in Italy

? Since the 1960s biologists have been trying to assess factors affecting cambial activity and latewood formation. A comparison of the phenology of the apical and lateral meristems could add new clues in order to understand mechanisms of the growth dynamic in conifers.

? This study compared needle and shoot growth with xylogenesis in Larix decidua, Pinus cembra and Picea abies during 2001 to verify if cambial resumption occurred after shoot and needle growth had begun, and if latewood initiation occurred when shoot lengthening was complete.

? In pine and spruce, needle and shoot lengthening was synchronous between mid-June and August while larches showed an early development of the needles at the end of May with a later shoot lengthening. Cell production lasted from mid-May to the beginning of August, with the first differentiating cells observed at the end of May, which indicated a earlier reactivation of the cambium with respect to needle and shoot growth.The first latewood cells began wall thickening between July and August, during the termination of needle and shoot lengthening.

? The hypothesis that cambial resumption occurs after shoot and needle elongation was rejected. The separation in time between apical growth and secondary wall thickening of latewood suggested the presence of an internal competition in resource allocation.

     

Tree diameter, height and stocking in even-aged forests

? Empirical observations suggest that in pure even-aged forests, the mean diameter of forest trees (D, diameter at breast height, 1.3 m above ground) tends to remain a constant proportion of stand height (H, average height of the largest trees in a stand) divided by the logarithm of stand density (N, number of trees per hectare): D = β(H ? 1.3)/ ln(N).

? Thinning causes a relatively small and temporary change in the slope β, the magnitude and duration of which depends on the nature of the thinning.

? This relationship may provide a robust predictor of growth in situations where scarce data and resources preclude more sophisticated modelling approaches.

     

Rank reversals in tree growth along tree size, competition and climatic gradients for four forest canopy dominant species in Central Spain

? Interspecific differences in tree growth patterns with respect to biotic and abiotic factors are key for understanding forest structure and dynamics, and predicting potential changes under climate change.

? Repeated observations from the Spanish Forest Inventory (SFI) were used to parameterize maximum likelihood estimators of tree growth as a function of tree size, competition indices and climate for Pinus pinaster, P. sylvestris, Quercus ilex and Q. pyrenaica.

? Significant responses to both biotic and abiotic factors were found, with interspecific differences in species performance along competition, temperature and precipitation gradients. Q. ilex was the species most tolerant to competition while P. pinaster was the species most sensitive to climatic variation. Species relative positions shifted along gradients of these factors with rank reversals in species performance along size, competition and climatic gradients.

? The results based on average growth matched previous forestry classifications and experimental studies on relative growth rate (RGR).

? When examining growth along studied abiotic and biotic gradients, a mismatch was found between species performance ranks as predicted by our models and information derived from previous knowledge. Those discrepancies highlight the relevance of ontogeny and environmental heterogeneity in defining species performance along competition gradients.

     

Effects of atmospheric and climate change at the timberline of the Central European Alps

? This review considers potential effects of atmospheric change and climate warming within the timberline ecotone of the Central European Alps. After focusing on the impacts of ozone (O₃) and rising atmospheric CO₂ concentration, effects of climate warming on the carbon and water balance of timberline trees and forests will be outlined towards conclusions about changes in tree growth and treeline dynamics.

? Presently, ambient ground-level O₃ concentrations do not exert crucial stress on adult conifers at the timberline of the Central European Alps. In response to elevated atmospheric CO₂ Larix decidua showed growth increase, whereas no such response was found in Pinus uncinata. Overall climate warming appears as the factor responsible for the observed growth stimulation of timberline trees.

? Increased seedling re-establishment in the Central European Alps however, resulted from invasion into potential habitats rather than upward migration due to climate change, although seedlings will only reach tree size upon successful coupling with the atmosphere and thus loosing the beneficial microclimate of low stature vegetation.

? In conclusion, future climate extremes are more likely than the gradual temperature increase to control treeline dynamics in the Central European Alps.

     

Radial distribution of quinones in plantation teak (Tectona grandis L.f.)

? Quinones are primarily responsible for the natural durability of teak.

? The previous paper in this series reported on the natural termite resistance of teak trees of different ages (8-, 30- and 51-year-old trees). In this study, the radial distribution of quinones (tectoquinone, lapachol, desoxylapachol and its isomer) and other components in the ethanol-benzene (1:2) extract were measured by means of gas chromatography.

? Significant differences in desoxylapachol or its isomer content were found among the outer heartwood of 8-, 30- and 51-year old trees, as well as between the inner and outer parts of the heartwood.

? All toxic quinone contents were positively correlated with the total extractive content. The highest correlation degree was measured in the isodesoxylapachol content.

? Although linearly related, only modest correlations were observed between the natural termite resistance parameters and the content of tectoquinone and isodesoxylapachol.