A statistical model for the prediction of the number of sapwood rings in Scots pine (Pinus sylvestris L.)

Dendrochronology is a well-established dating method for wooden objects, but due to surface processing of construction timber or natural degradation the dating of historical wood often relies on a prediction of the number of missing rings based on sapwood statistics. Since Scots pine (Pinus sylvestris L.) is one of the most common tree species in north-western Europe, the absence of reliable sapwood statistics and models for the prediction of missing sapwood rings for pine samples is remarkable. We have therefore produced sapwood statistics based on data from 776 pine trees with ages from 15 to 345 years. The material consists of both living trees and historical timber, with varying growth rates, geographic settings, and from different soil types. When the whole material is considered, the average age of the trees is 103 years, and the number of sapwood rings is 54 ± 15 (1 SD), but range from 18 to 129. Trees less than 100-years in age contained 46 ± 11 (1 SD) sapwood rings and had an average tree-ring width (TRW) of 1.76 mm. With increasing age, the average TRW decreased while the number of sapwood rings increased. The average TRW of 101–200-year-old trees is 0.99 mm while the samples contained 63 ± 12 (1 SD) sapwood rings. For trees older than 201 years, the average TRW is 0.64 mm while the number of sapwood rings increased to 85 ± 16 (1 SD). The two most important factors in determining the number of sapwood rings for a given tree when only heartwood statistics are available proved to be (i) the number of heartwood rings and (ii) the average TRW of the heartwood rings. For incomplete samples, we have therefore developed a statistical model based on the sample’s heartwood rings (number and average width) to compute a prediction interval for the total number of rings. The sapwood and heartwood statistics suggest a statistical model for the number of sapwood rings with mean that increase with the number of heartwood rings. Furthermore, the average number of sapwood rings decreases with the mean width of the heartwood rings. However, the predictive power of the mean width is limited when the number of heartwood rings has already been taken into account. Thus, we suggest making predictions for the number of sapwood rings using only the number of heartwood rings. Predictions of the number of sapwood rings based on the statistical model where convincing in the case of the three different datasets that were analysed. The certainty in these predictions was such that the width of the 80% and 95% prediction intervals ranged 28–34 and 45–52 sapwood rings, respectively. Additionally, we demonstrate how make predictions when there is information about the number of remaining sapwood rings in a given sample. To make the sapwood model available, we present a free online R package for fitting our models and an online software dashboard.     

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.     

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.     

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.     

Sapwood estimates of pedunculate oak (Quercus robur L.) in eastern Baltic

Pedunculate oak (Quercus robur L.) is one of the widely used and dendrochronologically investigated species in Europe. Still, it is a problematical dating object if its outermost section is missing partly or totally. Thus, we need sapwood estimation of living trees. As sapwood amount varies geographically, numbers of sapwood rings have been published for different regions in Europe but no such estimation has been done for the Baltic States yet. Therefore, this paper deals with the estimation of pedunculate oak sapwood growing in the eastern Baltic region, i.e. in Finland, Estonia, Latvia, and Lithuania.In total, 668 oak core samples of living trees from 43 stands were investigated. Ring widths were measured and the number of sapwood rings was determined according to two criteria: difference of colour and absence of tyloses in earlywood vessels. The samples were divided into two sets, according to the t_H-values between site chronologies and the major geobotanical sub-provinces. Thus, the nine Finnish and western Estonian sites were attributed to the western region and the 34 eastern Estonian, Latvian and Lithuanian sites to the eastern region.As the result of a statistical analysis, we explain that the number of oak sapwood rings ranges from 4.09 to 20.85 and 6.45 to 18.02 within 95% confidence limits in the western and eastern regions, respectively. For the three Baltic countries and southern Finland in general, we recommend to consider a sapwood estimate of 6.18–18.71 rings. Regarding earlier studies, the general European trend of decreasing sapwood ring number towards the east was confirmed. A geographical pattern of eastward decrease of the median sapwood ring number was noticed in the Baltics as well. The chronology based upon 668 samples of living oak trees from all sites covered the period of 1631–2008.     

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.     

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.     

Concentration gradients of free sterols,steryl esters and lipid phosphorus in the trunkwood of Scot's pine (Pinus sylvestris L.)

The amounts of free sterols, steryl esters and lipid phosphorus were determined in the sapwood and heartwood of mature, and in the outer and inner sapwood of young Pinus sylvestris trees. In the mature trees (up to 70 years old) the heartwood contains significantly higher amounts of free sterols than the sapwood. No radial gradient can be demonstrated in the amounts of steryl esters. Lipids extracted from the sapwood contain higher amounts of phosphorus than those from the heartwood. Stems of young Pinus sylvestris trees (up to 13 years old) show in the inner sapwood higher amounts of both free sterols and steryl esters than the peripheral younger wood zone. The inner sapwood of the young stems shows slightly higher amounts of lipid phosphorus than the outer sapwood. The results indicate that Pinus sylvestris accumulates both free sterols and steryl esters in the stems at a very early stage of the life cycle. Sterol accumulation in the innermost parts of the stems seems not to depend on heartwood formation.     

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.     

Respiration in the sapwood and heartwood of Robinia pseudoacacia

The respiratory activity of distinct sapwood and heartwood annual rings of the stem of Robinia pseudoacacia L. has been investigated. The oxygen uptake and the carbon dioxide release in the inner parts of the sapwood is enhanced in comparison with that in the outer parts. The heartwood rings have no measureable gas exchange.     

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.     

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.     

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.     

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.     

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.     

Size- and Age-dependent Variation in the Properties of Sap- and Heartwood in Black Locust (Robinia pseudoacaciaL.)

Dissection and mechanical bending experiments showed that the cross-sectional area and elastic moduli of sap- and heartwood varied within the trunk and branches as a function of the distance from the top of a 43-year-old black locust tree (Robinia pseudoacaciaL.). Wood in branches less than 1 m from the top of the tree consisted entirely of sapwood; the majority of the wood from more basipetal (and older) parts of the tree was heartwood. The Young's elastic moduli of sap- and heartwood increased towards the base of the trunk, and, on average, the modulus of the sapwood was 35%less than that of the heartwood. Younger, more distal tree limbs, therefore, were more flexible than older portions of the same tree. Simple bending experiments showed that the flexural rigidity of young limbs was governed by the location, physical properties, and the relative quantities of the two types of wood. The rigidity of limbs increased toward the base of the tree, and was dominated by sapwood in young limbs and by heartwood in the oldest parts of the tree. These trends predict that the younger, distal limbs of this tree can more easily deflect and bend in the wind, thereby reducing drag and the total bending moment on the tree trunk, while older limbs and the trunk are sufficiently rigid to support static self-loadings. Further study, however, is required to determine whether the trends reported here apply to all trees of this species and to trees of different species.     

人工长白落叶松树干边材、心材和树皮密度预测模型

基于黑龙江省林口林业局、东京城林业局和东北林业大学帽儿山实验林场的35株人工长白落叶松的解析样木数据,构建长白落叶松的边材、心材和树皮密度的Beta回归模型,采用赤池信息准则、决定系数、平均绝对偏差、均方根误差和似然比检验对模型的拟合优度进行比较评价,进而选取边材、心材和树皮密度的最优模型,最后采用刀切法对选择出的最优模型进行检验,评价模型预测能力。结果表明: 边材、心材和树皮密度的最优模型的自变量不完全相同,其中,边材密度与树木年龄、树高、相对高度和相对高度的平方关系较好,而心材密度最优模型的自变量为年生长量、相对高度和相对高度的平方,树皮密度最优模型的自变量为树木年龄、年生长量、相对高度和相对高度的平方。对最优模型分析可知,从树干基部到树梢,边材密度逐渐减小,心材密度先减小后增加,树皮密度先增加后减小。本研究所建立的Beta回归模型可以预估该研究区域的人工林内长白落叶松的边材、心材和树皮任意位置的木材密度,为树干平均密度和生物量的研究奠定基础。     

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.     

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.     

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