Radial and vertical variation of wood nutrients in Bornean tropical forest trees

Nitrogen, phosphorus, potassium, calcium, and magnesium concentrations in woody tissue are poorly documented, but are necessary for understanding whole-tree nutrient use and storage. Here, we report how wood macronutrient concentrations vary radially and along the length of a tree for 10 tropical tree species in Sabah, Malaysia. Bark nutrient concentrations were consistently high: 2.9–13.7 times greater than heartwood depending on the nutrient. In contrast, within the wood both the radial (sapwood vs. heartwood) and vertical (trunk bottom vs. trunk middle) variation was modest. Higher concentrations in sapwood relative to heartwood provide empirical support for wood nutrient resorption during sapwood senescence. Dipterocarp species showed resorption rates of 25.3 ± 7.1% (nitrogen), 62.7 ± 11.9% (phosphorus), and 56.2 ± 12.5% (potassium), respectively, while non-dipterocarp species showed no evidence of nutrient resorption in wood. This suggests that while dipterocarps have lower wood nutrient concentrations, this family is able to compensate for this by using wood nutrient resorption as an efficient nutrient conservation mechanism. In contrast to other nutrients, calcium and magnesium tended to accumulate in heartwood. Wood density (WD) showed little vertical variation along the trunk. Across the species (WD range of 0.33 to 0.94 mg/cm³), WD was negatively correlated with wood P and K concentration and positively correlated with wood Ca concentration. As our study showed exceptionally high nutrient concentrations in the bark, debarking and leaving the bark of the harvested trees on site during logging operations could substantially contribute to maintaining nutrients within forest ecosystems.     

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.     

Insights into intraspecific wood density variation and its relationship to growth,height and elevation in a treeline species

• The wood economics spectrum provides a general framework for interspecific trait–trait coordination across wide environmental gradients. Whether global patterns are mirrored within species constitutes a poorly explored subject. In this study, I first determined whether wood density co‐varies together with elevation, tree growth and height at the within‐species level. Second, I determined the variation of wood density in different stem parts (trunk, branch and twigs).
• In situ trunk sapwood, trunk heartwood, branch and twig densities, in addition to stem growth rates and tree height were determined in adult trees of Nothofagus pumilio at four elevations in five locations spanning 18° of latitude. Mixed effects models were fitted to test relationships among variables.
• The variation in wood density reported in this study was narrow (ca. 0.4–0.6 g cm^?3) relative to global density variation (ca. 0.3–1.0 g cm^?3). There was no significant relationship between stem growth rates and wood density. Furthermore, the elevation gradient did not alter the wood density of any stem part. Trunk sapwood density was negatively related to tree height. Twig density was higher than branch and trunk densities. Trunk heartwood density was always significantly higher than sapwood density.
• Negative across‐species trends found in the growth–wood density relationship may not emerge as the aggregate of parallel intraspecific patterns. Actually, trees with contrasting growth rates show similar wood density values. Tree height, which is tightly related to elevation, showed a negative relationship with sapwood density.

     

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.     

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.     

Resistance of Scots pine wood to Brown-rot fungi after long-term forest fertilization

The susceptibility of Scots pine (Pinus sylvestris L.) sap- and heartwood against the wood decaying brown-rot fungus (Coniophora puteana) was investigated after long-term forest fertilization at three different sites in central Finland. Different wood properties: wood extractives, wood chemistry, and wood anatomy were used to explain sap- and heartwood decay. Scots pine sapwood was more susceptible to decay than its heartwood. In one site, sapwood seemed to be more resistant to wood decay after forest fertilization whereas the susceptibility of heartwood increased. Significant changes in the sapwood chemistry were found between treatment and sites, however, no relationship between wood chemistry and wood decay was observed in the factor analysis. The results of this study show that there was an inconsistent relationship between decay susceptibility and fertilization and the measured physical and chemical attributes of the wood were not consistently correlated with the decay rate.     

The effect of water soluble Scots pine (Pinus sylvestris L.) and Sitka spruce [Picea sitchensis (Bong.) carr.] heartwood and sapwood extracts on the growth of selected Trichoderma species

Extractable materials from some timber species have been identified which prevent wood decay; however, little has been reported on the effect(s) of such materials against mould species that colonize timber. With increasing interest in the use of Trichoderma species, both as agents of permeability enhancement and biological control, more information is required on how chemical components within fresh and processed timber influence growth of Trichoderma. Fresh and dried samples of Scots pine and Sitka spruce sapwood and heartwood were leached in a Soxhlet apparatus and the resulting extract was combined with malt extract agar and inoculated with Trichoderma. Trichoderma isolates were inhibited to varying degrees by extractives removed from fresh and dried heartwood of the two timbers. Growth on sapwood extractives, however, showed a lesser degree of inhibition. The implications of the results for applications of Trichoderma in timber are discussed.     

Natural wood colouring process in Juglans sp. (J. nigra, J. regia and hybrid J. nigra 23 ×J. regia) depends on native phenolic compounds accumulated in the transition zone between sapwood and heartwood

 Radial distribution of soluble phenolics was investigated at different heights in stems of Juglans nigra, J. regia and hybrids J. nigra 23 × J. regia. Four major phenolic compounds were studied: hydrojuglone glucoside (HJG), quercitrin (QUER) and two unknown compounds characterized as two ellagic acid derivatives E1 and E2. HJG and E1 content increased gradually in the sapwood, peaked in the sapwood-heartwood transition zone, and decreased drastically in the heartwood. QUER was accumulated preferentially around the transition zone, and its content was relatively low in the outer part of the sapwood and in the inner part of the heartwood. E2 content was low in the sapwood and increased in the heartwood. The heartwood formation was marked by the accumulation of new soluble compounds. The relationship between wood extractives and wood colour were evaluated and discussed. HJG was considered to be a major precursor of heartwood colour providing chromophores through hydrolysis (deglucosylation), oxidation and polymerization processes. Received: 2 September 1997 / Accepted: 23 November 1997     

Formation of heartwood substances in the stemwood of Robinia pseudoacacia L. II. Distribution of nonstructural carbohydrates and wood extractives across the trunk

Summary The distributions of reserve carbohydrates and of three dominant heartwood extractives were determined in the trunkwood of Robinia pseudoacacia L. The trees were cut at different times of the year (September, November, January, and April). With the exception of the tree felled in January, all trunks exhibited highest contents of nonstructural storage carbohydrates (glucose, fructose, sucrose, and starch) in the youngest, outermost sapwood zone. With increasing depth of the trunk, the levels of carbohydrates decreased. At the sapwood-heartwood transition zone, only trace amounts of nonstructural carbohydrates were present. The heartwood itself contained no storage material. The wood zones of different ages of the trees cut in September, November, and January exhibited glucose/fructose ratios of approximately 1. In April, however, there was a shift to glucose. In the youngest sapwood the amounts of soluble sugars were higher in the earlythan in the latewood. Older zones of the sapwood and the sap-wood-heartwood transition zone showed the opposite behaviour. Three main wood extractives of Robinia were characterized and quantified: the flavanonol dihydrorobinetin (DHR), the flavonol robinetin (ROB) and a hydroxycinnamic acid derivative (HCA). Only DHR was present — in very low amounts — in the younger sapwood of all trunks investigated. Higher amounts (>1 mol/g dry weight) of this compound and the HCA were present in the sapwood-heartwood transition zone. DHR augmented within the heartwood up to a more or less constant level. HCA increased towards the heartwood and decreased again in the inner heartwood parts. ROB appeared in the innermost parts of the sapwood-heartwood transition zone and reached maximum values in older parts of the heart-wood. The results indicate that starch is hydrolyzed at the sapwood-heartwood boundary and thus represents a primary major source of hydroxycinnamic acid and flavonoid synthesis.Dedicated to Prof. Meinhart H. Zenk on the occasion of his 60th birthday     

Formation and properties of some wood extractives

Chemical and other data concerning the location and site of formation of heartwood extractives are discussed. The biological conditions in the transition zone adjacent to the heartwood boundary are briefly described and some properties of extractives are given. The classes of polyphenolic compounds found in the wood of Eucalyptus species are listed together with the details of ellagic acid derivatives which are the most common class in this genus.     

大花序桉心边材变异规律和候选基因挖掘研究

为探究大花序桉(Eucalyptus cloeziana)心材比例差异显著的不同家系间心边材变异规律,挖掘心材变异相关的候选基因,为珍贵用材树种高效培育及育种利用提供基因资源。以18 a生的2个心材比例差异显著的大花序桉家系为材料(家系1和2),各制作解析木3株,沿着树干以1 m为区间分段截取圆盘,测量东西和南北2个方向的带皮直径、去皮直径、总年轮数、边材年轮数、边材直径,并开展心材和边材径向和轴向分析。同时利用各解析木胸径处初生木质部样品进行DNA混池测序,发掘等位基因频率差异显著的SNP位点并挖掘相关功能基因。结果表明,大花序桉边材宽度和心材半径的方位变异中家系2大于家系1,平均差值分别为0.7和5.5 cm,在随树高的变异中,家系1和2的心材半径和心材年轮数的下降速率分别为0.40和0.64及0.43和0.36。两家系间基本密度差异显著,家系1为0.80～0.82 g/cm³,家系2为0.75～0.78 g/cm³。基本密度与树高、横截面半径和心材半径呈显著负相关,与顺纹抗拉强度、弦面硬度和部分力学性质呈显著正相关。利用DNA混池测序共...     

A Genomic and Molecular View of Wood Formation

Wood formation is a process derived from plant secondary growth. Different from primary growth, plant secondary growth is derived from cambium meristem cells in the vascular and cork cambia and leads to the girth increase of the plant trunk. In the secondary growth process, plants convert most of photosynthesized products into various biopolymers for use in the formation of woody tissues. This article summarizes the new developments of genomic and genetic characterization of wood formation in herbaceous model plant and tree plant systems. Genomic studies have categorized a collection of the genes for which expression is associated with secondary growth. During wood formation, the expression of many genes is regulated in a stage-specific manner. The function of many genes involved in wood biosyntheses and xylem differentiation has been characterized. Although great progress has been achieved in the molecular and genomic understanding of plant secondary growth in recent years, the profound genetic mechanisms underlying this plant development remain to be investigated. Completion of the first tree genome sequence (Populus genome) provides a valuable genomic resource for characterization of plant secondary growth.     

Effect of Different Positions of Bordered Pit Torus in Yezo Spruce on Its Permeability

The green wood of yezo species (Picea jezoensis var. komarovii (V. Vassil.) Cheng et L. K. Fu) was treated with air drying and ethanol exchange drying and by soaking the air-dried wood in water followed by ethanol change drying. The results showed that the proportion of aspirated pits in the respective earlywood and latewood of the air-dried sapwood was increased to 99% and 81%, while that of heartwood was also as high as 97% and 86% because most of the bordered pits were aspirated at its green state. The permeability of air-dried sapwood and heartwood was as low as 0.114 and 0.045 darcy respectively. The proportion of aspirated pits in the earlywood and latewood of sapwood after ethanol exchange drying was very low (8% and 17% respectively), whereas that of heartwood was very high (97% and 86% respectively) since most of the pits in it were aspirated at its green state, so the ethanol exchange drying failed to prevent pit aspiration. The permeability of sapwood and heartwood after ethanol exchange drying was 11.713 and 0.074 darcy respectivly, which was increased 101.5 times and 62.0% over the permeability at air-dried state. t-test showed that the difference of permeability of ethanol exchange drying versus air drying for sapwood was very significant at the level of 0.1%, but was not significant for heartwood. The proportion of aspirated pits in earlywood and latewood of air-dried sapwood and heartwood after soaking in water followed by ethanol exchange drying was decreased by 18% and 22%, and 0 and 17%, respectively, while the permeability of sapwood and heartwood was 0.439 and 0.060 darcy respectively, which was increased by 85% and 49% respectively. The permeability difference of soaking sapwood and heartwood versus their controls was very significant at the 0.1% level by t-test.     

Mechanical Properties of Black Locust (Robinia pseudoacacia L.) Wood. Size- and Age-dependent Variations in Sap- and Heartwood

Variations in the density and stiffness (Young's elastic modulus)of fresh wood samples drawn from different parts of the threemain trunks of a 32-year-old black locust tree,Robinia pseudoacacia(measuring 19.8 m at its highest point), were studied to determinewhether tree ontogeny can achieve a constant safety factor againstmechanical failure. Based on the properties of isolated woodsamples, the fresh density of sapwood decreased along radialtransects from bark to pith, while that of progressively olderheartwood samples increased, on average, towards the centreof each of the three trunks. Along the same radial transects,the Young's elastic modulus of sap- and heartwood increased.In terms of longitudinal changes in wood properties, mean woodmoduli (averages of sap- and heartwood samples) increased, onaverage, towards the base of each of the three trunks of thetree. However, the mean fresh densities of wood samples increasedtowards the top and the bottom of each trunk and were lowestroughly near trunk mid-length. The mean density-specific stiffness(the quotient of Young's modulus and fresh density) of woodwas thus lower toward the top and the bottom of the trunks andhighest near trunk mid-length. Mean values of fresh wood density-specificstiffness were used to estimate the critical buckling heightsfor sections of the trunks differing in diameter and age. Theseestimates indicated that ontogenetic variation in the physicalproperties and relative amounts of sap- and heartwood in trunkscould maintain a constant factor of safety (approximately equalto 2) as a sapling grows in height and girth into a mature tree.This expectation was supported by data from 16 black locusttrees differing in height and diameter at breast height (DBH). Wood; elastic properties; tree height; biomechanics