Sapwood characteristics of Quercus robur species from the south-western part of the Pannonian Basin

We analysed sapwood characteristics in 344 pedunculate oak (Quercus robur L.) samples from the south-western part of the Pannonian Basin. The samples came from 13 sites, located in Slovenia, Croatia and Serbia. The trees had an average of 13.3 sapwood rings, with a minimum of 5 and maximum of 32. Fifteen log-log linear regression models were employed to assess the statistical relationship between sapwood and heartwood variables. The number of sapwood rings (N_SW), which is usually needed in dendroarchaeological dating, is significantly related to the number of heartwood rings (N_HW), heartwood width (W_HW) and heartwood growth rate (G_HW). Older and more slowly growing trees had a higher average number of sapwood rings. Using N_HW and W_HW, we employed an additional multiple regression model and calculated coefficients for N_SW predictions for real-world dendroarchaeological dating from the south-western part of the Pannonian Basin.     

Heartwood and sapwood development within maritime pine (<Emphasis Type="Italic">Pinus pinaster </Emphasis>Ait.) stems

Heartwood and sapwood development in maritime pine (Pinus pinaster Ait.) is reported based on 35 trees randomly sampled in four sites in Portugal. It was possible to model the number of heartwood rings with cambial age. The heartwood initiation age was estimated to be 13 years and the rate of sapwood transformation into heartwood was 0.5 and 0.7 rings year^–1 for ages below and above 55 years, respectively. Reconstruction of heartwood volume inside the tree stem was made by visual identification by image analysis in longitudinal boards along the sawn surfaces. This volume was integrated into the 3D models of logs and stems developed for this species representing the external shape and internal knots. Heartwood either follows the stem profile or shows a maximum value at 3.8 m in height, on average, while sapwood width is greater at the stem base and after 3 m remains almost constant up the stem. Up to 50% of tree height heartwood represents 17% of stem volume, in 83-year-old trees and 12–13% in 42 to 55-year-old trees. Tree variables such as stem diameter, DBH and tree total height were found to correlate significantly with the heartwood content.     

Estimating missing sapwood rings in three European gymnosperm species by the heartwood age rule

Precise dating of the year of felling is one intended outcome of dendrochronology. However, occasionally some or all sapwood rings might be missing, either due to deterioration or because they were carved off, or for some other reason. Consequently, while heartwood is preserved, sapwood might be fully or partially missing. In such cases, the year of felling must be estimated by adding a suitable number of sapwood rings. A heartwood age rule (HAR) has been advocated for Scots pine and adapted to European larch and Cembra pine, implying a linear relationship between sapwood ring count and the square root of heartwood ring count, largely irrespective of position in the stem. The same rule applied to all observations of a species, irrespective of silviculture, location or fertility of the growth site. Scots pine had twice or thrice as many sapwood rings as Cembra pine, which had 10% more rings than larch. The magnitude of model residuals was proportional to estimated sapwood ring count. Relative residuals were roughly normally distributed. To be applicable in Bayesian modelling in dendrochronology analyses, detailed information on model errors has been provided.     

The Zurich method for sapwood estimation

We present a new predictor for the estimation of the number of missing sapwood rings in oak. It circumvents a number of problems with predictors used in traditional sapwood estimation procedures such as mean ring width or tree age. Instead, we use the mean ring width of the last 9 heartwood rings and the first sapwood ring, which mirrors the growth level during sapwood formation. We present a model to produce a 95 % prediction interval for the felling year. Our method accounts for the skewed distribution of the number of sapwood rings towards higher values.     

Dendroclimatic signals in the pine and spruce chronologies in the Solovetsky Archipelago

Searching for a robust tree-ring parameter useful for paleoclimatic purposes is one of the most demanding topics in the modern paleoscience. Since Blue Intensity has already expressed itself in different geographical locations all over the world as a possible replacement for maximum density, close attention is paid to investigate features of the inferred signal. The Solovki Islands is a unique location in Northern Russia where two important factors that make this territory attractive for developing a long tree-ring chronology have been met: modern long-living trees and building activities using old trees that were started by monks in the middle of the 16th century. The main goal of the research is to develop pine and spruce chronologies based on tree-ring width (TRW) and delta Blue Intensity (dBI) and to assess the ability of these parameters to be used as climate predictors. As a result, 14 conifer chronologies from 7 sites (4 for pine and 3 for spruce) were developed. The composite pine and spruce chronologies span a period of 474 and 378 years each. Cross-correlation of dBI-based chronologies of both conifers is high (r = up to 0.71 while for TRW-based chronologies it is lower on average (−0.18 to 0.63). Intra-species correlation of TRW chronologies in some cases achieved even negative values (r = −0.18. Discrepancies found between TRW chronologies of pine and spruce could be explained by differences in climatic signals. Response function analysis with monthly temperatures revealed that growth of pine depends on the previous August, while spruce has a temporally stable and strong relation to June temperatures. Compared to TRW, dBI-based chronologies have a high correlation with summer temperatures (r = 0.64 and 0.66 for spruce and pine, respectively). Presented research points out the importance of the response function analysis suggesting that depending on goals of the study several tree-ring parameters could be used, e.g., tree-ring width of spruce responses to June temperatures, while dBI to the whole summer.     

From desert to rainforest, sapwood width is similar in the widespread conifer Callitris columellaris

The process that transforms conductive sapwood to non-conductive heartwood in trees is poorly understood. Here, we use natural variation in climate to examine the environmental control of sapwood width in a widespread conifer species. We hypothesised that if sapwood width is linked to transpirational load, there would be a positive association between sapwood width, and continental gradients in mean annual rainfall, whereas age-related conversion to heartwood would be revealed from estimates of the age of the inner-most sapwood ring. Using the widespread Australian conifer Callitris columellaris we took cores from trees at 85 sites spanning a range of 168–2,117 mm in mean annual rainfall, and 14–28 °C in mean annual temperature. We found that sapwood width was remarkably similar throughout the species range, being only slightly lower in the tropics than the arid or temperate zone. There was a weak negative relationship between sapwood width and mean annual rainfall, which is in the opposite direction expected from transpirational control of sapwood width. Sapwood growth rings were wider, but there were fewer of them in the tropics than elsewhere, indicating conversion to heartwood occurred earlier here. Together with an earlier finding that tracheid diameter was largest in the tropics, our results show that differences amongst climate zones more strongly influence the hydraulic properties of sapwood than its amount.     

Variation of heartwood and sapwood in 18-year-old Eucalyptus globulus trees grown with different spacings

Heartwood and sapwood development was studied in 18-year-old Eucalyptus globulus trees from pulpwood plantations with different spacings (3 × 2, 3 × 3, 4 × 3, 4 × 4 and 4 × 5 m), on cross-sectional discs taken at breast height. The trees possessed a large proportion of heartwood, on average 60% of the wood cross-sectional surface. Spacing was a statistically significant source of variation of heartwood area, which ranged between 99 and 206 cm² for the closer (3 × 2) and wider (4 × 5) spacings, respectively. There was a positive and high statistical significant correlation between heartwood diameter and tree diameter (heartwood diameter = −0.272 + 0.616 dbh; r ² = 0.77; P < 0.001), and larger trees contained more heartwood regardless of spacing. Heartwood proportion in cross-section remained practically constant between spacings but increased with tree diameter class: 55.1, 62.2, 65.0 and 69.5% for diameter at breast height classes <15, 15–20, 20–25 and >25 cm, respectively. The sapwood width did not depend on tree diameter growth and remained practically constant at an average of 18 mm (range 15–21 mm), but sapwood area showed a good linear regression with tree diameter. Therefore, tree growth enhancement factors, such as wide spacings, will induce formation of larger heartwoods that can negatively impact raw-material quality for pulping. The increase in heartwood in relation with tree dimensions should therefore be taken into account when designing forest management guidelines.     

Growth rate and ring width variability of teak, Tectona grandis (Verbenaceae) in an unmanaged forest in East Timor

Teak (Tectona grandis) is one of the most valuable timbers in international trade and an important species for tropical forestry. Teak is found on the island of East Timor but no information is available on teak growth from this region. A pure stand planted in 1940-50 in the North of East Timor and left unmanaged was studied. Fifteen trees were sampled in October-November 2003 and stem discs taken at three height levels of its height (1.7m, 9.5m and 18.7m), and cores were collected at DBH. Transverse surfaces of the discs and cores were polished for ring identification. Core cross sections were first digitized and disc cross sections were observed under the microscope. Three randomly selected radii were analyzed in each disc. Ring width measurement and ring counting were done using image analysis software. The distinction between heartwood and sapwood was performed macroscopically by colour difference, and heartwood radius and sapwood width were measured. The relationship between stem and heartwood radius was studied for each disc and heartwood percentage by radius was determined. Radial ring width curves are presented for the different axial positions within the stem, and ring width variability was analyzed. Growth rates were calculated and age-radius relationships were estimated using cumulative growth curves. Growth rings were large and well defined in the juvenile phase, reflecting the specie's fast-growing character. The year-to-year variation of ring width showed a similar pattern among trees. Mean ring width ranged between 4.3-7.3mm for the first 20 years and 3.3-5.1mm for 30 to 45 years. Pith eccentricity was evident in the lower part of the stem and ring wedging occurred. On average, heartwood represented 84% of the radius and sapwood contained 6 to 11 rings. The age-related variation of ring width and the occurrence in the lower part of the tree stems of eccentricity and wedging rings, highlights the importance of appropriate stand management, particularly regarding basal density distribution over time, whenever optimized timber production is envisaged.     

Diversity and decay ability of basidiomycetes isolated from lodgepole pines killed by the mountain pine beetle

When lodgepole pines (Pinus contorta Douglas ex Louden var. latifolia Engelm. ex S. Watson) that are killed by the mountain pine beetle (Dendroctonus ponderosae) and its fungal associates are not harvested, fungal decay can affect wood and fibre properties. Ophiostomatoids stain sapwood but do not affect the structural properties of wood. In contrast, white or brown decay basidiomycetes degrade wood. We isolated both staining and decay fungi from 300 lodgepole pine trees killed by mountain pine beetle at green, red, and grey stages at 10 sites across British Columbia. We retained 224 basidiomycete isolates that we classified into 34 species using morphological and physiological characteristics and rDNA large subunit sequences. The number of basidiomycete species varied from 4 to 14 species per site. We assessed the ability of these fungi to degrade both pine sapwood and heartwood using the soil jar decay test. The highest wood mass losses for both sapwood and heartwood were measured for the brown rot species Fomitopsis pinicola and the white rot Metulodontia and Ganoderma species. The sap rot species Trichaptum abietinum was more damaging for sapwood than for heartwood. A number of species caused more than 50% wood mass losses after 12 weeks at room temperature, suggesting that beetle-killed trees can rapidly lose market value due to degradation of wood structural components.     

Relationship of heartwood traits with diameter growth, implications for genetic selection in Pinus radiata

The heartwood of radiata pine (Pinus radiata D. Don) is not favoured commercially because of its darker appearance, unreliable durability and difficulty to treat chemically; consequently, light-coloured sapwood, which is more easily treated with decay-resistant chemicals, is of more value. To decrease the amount of heartwood or increase sapwood cross-sectional area by genetic selection, the genetics of sapwood cross-sectional area (SWA), sapwood area percentage (SWAP) and number of heartwood rings (HWR) is of interest. We investigated genetic variation in these three variables from increment cores from trees from two trial series, each planted at two sites, from a total of some 330 families (189 were half-sib and 142 were full-sib) with 10 to 30 individuals assessed per family. Trials were assessed between 16 and 23 years old when all trees had started to develop heartwood. The three sapwood-related traits were moderately heritable across pairs of sites ( h ^ i 2 $ {\widehat{h}}_i^2 $ = 0.2 to 0.4) indicating opportunity for selection. SWA was strongly correlated (r _a ?=?0.9) with diameter. SWAP and HWR were also strongly correlated (r _a ?=??0.9). The number of HWR was independent of tree diameter, providing the opportunity through genetic selection to reduce heartwood area whilst increasing tree growth rate.     

Effects of age and site quality on the distribution of biomass in Scots pine (Pinus sylvestris L.)

The distribution of the above-ground and below-ground biomass of Scots pine in southern Finland were investigated in trees of different ages (18–212 years) from two types of growth site. Secondly, some structural regularities were tested for their independence of age and growth site. Trees were sampled from dominant trees which could be expected to have a comparable position in stands of all ages. All stands were on sorted sediments. The biomass of the sample trees (18 trees) was divided into needles, branch sapwood and heartwood, stem sapwood and heartwood, stem bark, stump, large roots (diameter >20 cm), coarse roots (five classes) and fine roots. The amount of sapwood and heartwood was also estimated from the below-ground compartments. Trees on both types of growth site followed the same pattern of development of the relative shares of biomass compartments, although the growth rates were faster on the more fertile site. The relative amount of sapwood peaked after canopy closure, coinciding with the start of considerable heartwood accumulation. The relative amount of needles and fine roots decreased with age. The same was true of branches but to a lesser degree. The relative share of the below-ground section was independent of tree age. Foliage biomass and sapwood cross-sectional area were linearly correlated, but there were differences between the growth sites. Needle biomass was linearly correlated with crown surface area. The fine root to foliage biomass ratio showed an increasing trend with tree age.     

Cellular level observation of water loss and the refilling of tracheids in the xylem of Cryptomeria japonica during heartwood formation

The mechanism of heartwood formation in Cryptomeria japonica D. Don has long been studied since heartwood formation is a fundamental physiological feature of trees. In this study, the water distribution in the xylem of C. japonica was investigated at the cellular level to reveal the role of water distribution in the xylem during heartwood formation. Samples were taken from different heights of each trunk, in which the phases of heartwood formation differed. These were designated as SIH, which consisted of sapwood, intermediate wood, and heartwood; SI, which consisted of sapwood and intermediate wood but no heartwood; and S-all, which consisted entirely of sapwood. Cryo-scanning electron microscopic observations of the heartwood-formed (SIH) and non-heartwood-formed (SI and S-all) xylem revealed different patterns of water distribution changes in tracheids between the latewood and earlywood. In the latewood, almost all tracheids were filled with water in all areas from the sapwood to the heartwood (98–100% of tracheids had water in their lumina). In the earlywood, however, the water distribution differed between the sapwood (95–99%), intermediate wood (7–12%), and heartwood (4–100%). Many of the tracheids in the xylem, where the sapwood changed to intermediate wood lost water. In the heartwood, some tracheids remained empty, while others were refilled with water. These results suggest that the water distribution changes in individual tracheids are closely related to heartwood formation. Water loss from tracheids may be an important factor inducing heartwood formation in the xylem of C. japonica.     

Development of heartwood in response to water stress for radiata pine in Southern New South Wales, Australia

Heartwood development and other functional changes in stem conductance in response to water stress in radiata pine were investigated using two contrasting climatic areas (high-altitude sub-alpine vs. warm–dry inland) of the Hume region of New South Wales, Australia. The study included mature (34.5–36.5 years old) and young stands (10–11 years old) measured under normal climate and during an extreme drought. The effect of water stress on heartwood development was examined using sapwood percentage, sapwood saturation, development of dry sapwood and evidence of cavitation in sapwood. Trees at the warm–dry site developed heartwood at faster rates than on the high-altitude site. At breast height, the mature stands of the warm–dry site had 8–14 % less sapwood. Extensive cavitation towards the sapwood/heartwood boundary occurred in some of the mature and young stands on the warm–dry site. We postulated that in water-limiting environments, cavitation of the inner sapwood precedes heartwood formation and is an adaptation mechanism that regulates stem conductance capacity and thus water use in the tree. The drought of 2006 led to decreases in moisture associated with cavitation not previously reported for radiata pine and demonstrated the drought hardiness of the species. In the warm–dry site, breast-height sapwood saturation dropped to 58 and 82 % for suppressed and average-sized trees in a mature unthinned stand; and 75–78 % for two young stands. These saturation levels, however, only imply average values as some cells cavitated whilst others were fully saturated. Cavitation occurred in a localized fashion affecting small to large groups of cells.     

Xylem water content and wood density in spruce and oak trees detected by high-resolution computed tomography

Elucidation of the mechanisms involved in long-distance water transport in trees requires knowledge of the water distribution within the sapwood and heartwood of the stem as well as of the earlywood and latewood of an annual ring. X-ray computed tomography is a powerful tool for measuring density distributions and water contents in the xylem with high spatial resolution. Ten- to 20-year-old spruce (Picea abies L. KARST.) and oak (Quercus robur) trees grown in the field were used throughout the experiments. Stem and branch discs were collected from different tree heights, immediately deep frozen, and used for the tomographic determinations of spatial water distributions. Results are presented for single-tree individuals, demonstrating heartwood and sapwood distribution throughout their entire length as well as the water relations in single annual rings of both types of wood. Tree rings of the sapwood show steep water gradients from latewood to earlywood, whereas those of the heartwood reflect water deficiency in both species. Although only the latest two annual rings of the ringporous species are generally assumed to transport water, we found similar amounts of water and no tyloses in all rings of the oak sapwood, which indicates that at least water storage is important in the whole sapwood.     

Seasonal changes of pinosylvin distribution in the sapwood/heartwood boundary of Pinus sylvestris

Changes in pinosylvin concentration and distribution across the sapwood/heartwood boundary were studied on Scots pine (Pinus sylvestris L.) tree stems to detect seasonal activity in heartwood formation. Pinosylvin concentrations were measured with FT-(NIR) Raman spectroscopy for a total of 96 trees equally distributed on 16 different sampling occasions. In another experiment, cores were sampled every month from six individual Scots pine trees from June to October and analysed for pinosylvin. There was a great between-tree variation in concentration and spatial distribution of pinosylvin. No seasonal trend in the distribution pattern or concentration of pinosylvin was found. Therefore, the results indicate that there is no specific period during the year when heartwood is formed. Received: 12 February 1998 / Accepted: 3 May 1999     

Towards a new approach for dendroprovenancing pines in the Mediterranean Iberian Peninsula

Dendroprovenancing studies frequently use site chronologies to identify the origin of archaeological and historical timber. However, radial growth (tree-ring width, TRW) of tree species is influenced by both local and regional climate scales. Here we investigate how the use of annually-resolved Blue Intensity (BI) measurements can enhance dendroprovenancing precision of black pine (Pinus nigra Arn.) and Scots pine (P. sylvestris L.) on the Iberian Peninsula. Principal Component Gradient Analyses (PCGA) was used to assess geographical patterns of annual variation in different TRW and BI proxies of pine trees from two mountain ranges in the Central System and Andalusia in Spain. Local climate-growth relationships were quantified to identify underlying causes of identified groups with diverse growth patterns. Two distinct elevational groups were observed when performing PCGA on latewood BI time series with the response to summer drought as the main factor causing the differences. Both P. nigra and P. sylvestris BI time series were found to be more related to summer drought at low-elevation sites showing an increase in sensitivity at lower latitudes. PCGA of TRW time series allowed to discriminate between trees from Andalusia and Central System within the elevation groups. February and October temperatures were found to be the main climatic factors causing the differences in TRW time series among the high- elevation sites, whereas for low-elevation trees it was the average winter temperature influencing TRW. A subsequent leave-one-out analyses confirmed that including latewood BI time series improves the precision of dendroprovenancing of pine wood in the Iberian Peninsula.     

Irregular Heartwood Formation in Maritime Pine (Pinus pinaster Ait):Consequences for Biomechanical and Hydraulic Tree Functioning

As the proportion of sapwood (SW) transformed into heartwood(HW) is irregular both radially and longitudinally in trunksof Maritime pine (Pinus pinaster Ait.), it has been suggestedthat HW formation is a developmental process, regulated internallywithin the tree. In trees where stem growth is eccentric dueto stem lean or wind action, the number of annual growth ringsof SW transformed into HW is greater on the compressed sideof the tree. To determine the contribution to bending stiffness,if any, of this prematurely formed HW, four point bending testswere carried out on fresh HW and SW samples taken from the samegrowth ring, or neighbouring growth rings, at different cross-sectionalpositions at a height of 2 m from six 52-year-old Maritime pines.The mean (±s.e.) modulus of elasticity (E) was 7.6 ±0.3 GPa (longitudinal direction) for all samples. No significantdifferences in E were found between HW and SW; thus HW doesnot play a significant mechanical role in bending stiffness.To test a second hypothesis that early HW formation on the compressedside of trees may maintain a constant, optimal volume of SWaround the tree, the Pipe Model Theory was applied to 12 52-year-oldleaning Maritime pines (angle of lean varied from 0–22°).The surface area (S) of the SW was determined at different heightsup the trunk and correlated with crown surface area (S_crown).Regressions between S_SWandS_crown were highly significant, therebysupporting the theory that HW formation and extension is controlledinternally in Maritime pine. HW formation in Maritime pine thenserves to maintain an optimal proportion of functional SW whichis an important criterion for survival in a species often subjectedto severe drought for long periods. Copyright 2001 Annals ofBotany Company Heartwood, sapwood, bending test, Pipe Model Theory, Pinus pinaster     

Effects of Infestation by the Balsam Woolly Aphid, Adelges Piceae (Ratz.) on the Ultrastructure of Bodered-Pit Membranes of Grand Fir, Abies grandis (Doug.) Lindl.

The ultrastructure of bordered-pit membranes in normal grandfir trees, Abies grandis (Doug.) Lindl., was compared with thatin trees infested with the blasam woolly aphid, Adelges piceae(Ratz). In sapwood of non-infested trees the membranes of earlywoodpits were well perforated, whereas those of latewood pits wereeither heavily incrusted or incompletely developed and showedfew perforations. In the heartwood pit membranes from both earlywoodand latewood were heavily incrusted. In aphid-infested trees all the pit membranes from the sapwoodwere incrusted and resembled those from heartwood of non-infestedtrees. These incurstations reduced the number of pores in themargo of pit membranes, and could account for the reduced permeabilityto water reported for sapwood trees attacked by the aphid. Wesuggest that the incrustation of pit membranes in sapwood inAbies grandis infested with Adelges piceae occurs because theseaphids cause heartwood to form prematurely.     

Food reserves of scots pine (Pinus sylvestris L.)

Summary Starch, soluble sugars, triacylglycerols, diacylglycerols and free fatty acids were measured in 30-year-old Scots pine (Pinus sylvestris L.) trees during an annual cycle in the sapwood (youngest ten xylem rings). The radial distribution of carbohydrates and lipids was studied in the trunkwood of 90 -to 150-year-old Scots pine trees collected at the end of the growing season. Determination of the compounds was performed using specific enzymatic assays, capillary gas chromatography and thin layer chromatography. The amounts of glucose, fructose, sucrose, and galactose/arabinose in the sapwood were slightly higher in winter than in summer. Raffinose/stachyose increased up to 5-fold during the cold period. At the beginning of the growing season starch amounts rose, and then decreased in summer and autumn. No concentration changes were observed in the total amounts of diacylglycerols and fatty acids throughout the year. Triacylglycerol levels were slightly higher in February than in summer and autumn. Relative frequencies of individual fatty acids were similar in all lipid fractions. Glucose, fructose, sucrose, starch and triacylglycerols disappeared almost entirely at the transition zone from sapwood to heartwood. In contrast, free fatty acids and galactose/arabinose rose in centripetal direction, and diacylglycerols remained constant across trunk cross-sections. The relative amounts of individual fatty acids changed markedly in the free fatty acid fraction and in the triacylglycerols when crossing the sapwood-heartwood boundary. Concentration changes of reserve materials are discussed in relation to season, mobilization and translocation processes, dormancy, frost resistance, and heartwood formation. The results are compared to those found in needles.     

Formation and vertical distribution of sapwood and heartwood in Cryptomeria japonica D. Don

The formation and vertical distribution of sapwood and heartwood were studied with a 45-year-old Cryptomeria japonica D. Don. The tree was grown at a plantation with 1.5 m × 3.0 m spacing near Miao-Li, Taiwan and was felled on 27 February 1992. The thickness of sapwood and heartwood was expressed by a ring count and a linear measurement. The east-west (E-W) wood strips were collected from 0.3 m above ground upwards to the top of the tree at 2.5 m intervals. The sapwood thicknesses from the base to the 10.3 m tree level height are around 20–22 growth rings and 42±2 mm. At the top of the tree, the sapwood thickness is narrower. The heartwood, which decreases in thickness with increasing tree level heights is not found at the top of the tree. The heartwood appears as a conical shape in the tree trunk. There is no statistical difference in sapwood/heartwood thickness between E-W aspects. Tree level heights and the tree level age were found to be important parameters in determining the thickness of sapwood/heartwood.