Radial wood allocation in Schizolobium parahyba

? Premise of the study: Pioneer species of tropical trees allocate wood specific gravity (SG) differently across the radius. Some species exhibit relatively uniform, low SG wood, whereas many others exhibit linear increases in SG across the radius. Here, we measured changes in SG across the radius of Schizolobium parahyba (Fabaceae-Caesalpinioideae), a wide-ranging, neotropical pioneer, used extensively in land reclamation and forest restoration in Brazil. ? Methods: Pith-to-bark radial wood cores were extracted with increment borers from 42 trees at five sites, in Central and South America. Cores were cut into 1-cm segments whose specific gravities were determined and analyzed via linear and nonlinear regression. Wood specific gravity, very low initially at 0.15-0.20, doubled or tripled across the tree radius to 0.45-0.65 for large adults. ? Key results: Unlike linear increases in other tropical pioneers, the increases in Schizolobium were nonlinear (convex up). At one site with even-aged trees, the magnitude of the radial increase was similar in all trees, despite a 4-fold difference in diameter among trees, implying that the radial increases in Schizolobium were regulated by tree age, not by tree size. ? Conclusions: This unique pattern of development should provide an extended period of growth when SG is low, facilitating hyper-extension of the bole, at some risk of structural failure. Later in growth, the SG rate of increase accelerates, reinforcing what was a precarious bole. Overall, these results suggest a third model for xylem allocation in tropical trees, a model that may be associated with monopodial stem development and limited life span.     

Does canopy position affect wood specific gravity in temperate forest trees?

The radial increases in wood specific gravity known in many tree species have been interpreted as providing mechanical support in response to the stresses associated with wind loading. This interpretation leads to the hypothesis that individuals reaching the canopy should (1) be more likely to have radial increases in specific gravity and (2) exhibit greater increases than individuals in the subcanopy. Wood specific gravity was determined for three species of forest trees (Acer rubrum, Fagus grandifolia and Tsuga canadensis) growing in central Massachusetts, USA. Acer rubrum shows radial increases in specific gravity, but these increases are not more pronounced in canopy trees; the other two species show a pattern of radial decreases. The degree of radial increase or decrease is influenced by tree height and diameter. Of the dominant tree species for which we have data, A. rubrum, Betula papyrifera and Pinus strobus show radial increases in specific gravity, whereas F. grandifolia, T. canadensis and Quercus rubra show decreases. The occurrence of radial increases in B. papyrifera and P. strobus, which are often canopy emergents, suggests that it is overall adaptive strategy that is important rather than position (canopy vs. subcanopy) of any individual tree. It is suggested that radial increases in specific gravity are associated with early-successional status or characteristics and decreases with late-successional status or persistence in mature forest.     

Wood density and its radial variation in six canopy tree species differing in shade-tolerance in western Thailand

Background and Aims

Wood density is a key variable for understanding life history strategies in tropical trees. Differences in wood density and its radial variation were related to the shade-tolerance of six canopy tree species in seasonally dry tropical forest in Thailand. In addition, using tree ring measurements, the influence of tree size, age and annual increment on radial density gradients was analysed.

Methods

Wood density was determined from tree cores using X-ray densitometry. X-ray films were digitized and images were measured, resulting in a continuous density profile for each sample. Mixed models were then developed to analyse differences in average wood density and in radial gradients in density among the six tree species, as well as the effects of tree age, size and annual increment on radial increases in Melia azedarach.

Key Results

Average wood density generally reflected differences in shade-tolerance, varying by nearly a factor of two. Radial gradients occurred in all species, ranging from an increase of (approx. 70%) in the shade-intolerant Melia azedarach to a decrease of approx. 13% in the shade-tolerant Neolitsea obtusifolia, but the slopes of radial gradients were generally unrelated to shade-tolerance. For Melia azedarach, radial increases were most-parsimoniously explained by log-transformed tree age and annual increment rather than by tree size.

Conclusions

The results indicate that average wood density generally reflects differences in shade-tolerance in seasonally dry tropical forests; however, inferences based on wood density alone are potentially misleading for species with complex life histories. In addition, the findings suggest that a ‘whole-tree’ view of life history and biomechanics is important for understanding patterns of radial variation in wood density. Finally, accounting for wood density gradients is likely to improve the accuracy of estimates of stem biomass and carbon in tropical trees.Key words: Radial gradients, shade-tolerance, tree biomass estimates, tree rings, tropical trees, wood density     

Wood specific gravity and its radial variations: the many ways to make a tree

In a study of radial trends in specific gravity of ~100 individuals of six mixed-northern-hardwood-forest tree species, three species (Acer rubrum, Pinus strobus, and Betula papyrifera) showed radial increases and three species (Quercus rubrum, Tsuga canadensis, and Fagus grandifolia) radial decreases. Analysis of these data, together with a larger data set including both temperate and tropical tree species, focuses on relationship of radial trends to successional status, variation in inside versus outside wood, and factors affecting degree and direction of radial trend. We propose a model for radial trends in which (1) radial increases associated with low values of specific gravity are an early-successional characteristic and radial decreases associated with high values of specific gravity late successional, (2) the differences in specific gravity and its radial trends reflect growth strategy and biomechanical considerations, and (3) the convergence in values seen in older trees is due to constraints relating to support.     

The Buttressed Blue Marble Tree: Wood and Growth Characteristics of Elaeocarpus angustifolius (Elaeocarpaceae)

Elaeocarpus angustifolius, a forest tree native to Australia,was introduced into the Hawaiian Islands and is now naturalizedlocally. The main purpose of this study was to test the hypothesisthat a radial increase in wood specific gravity was presentin trunk wood of these trees, which grow quite large (diameterat breast height=200 cm) and have massive buttress systems.Information on buttress height and number and specific gravityof the outer trunk wood (sampled at breast height), as wellas anatomical characteristics pertaining to conduction (vesseldiameter and density), was obtained from a range of different-sizedtrees. Both buttress height and number increase with increasingtree diameter. Wood specific gravity has a median value of 0.49and increases more than 50% over the range of tree diametersstudied. Vessel diameter increases over two-fold and vesselfrequency decreases with increasing tree diameter, althoughvery large trees (diameters>70 cm) exhibit more variability.Trees have buttresses spaced evenly around the circumference(maximum of 15–20) and exhibit no difference in wood specificgravity on the leeward and windward sides in spite of theirlocation in the trade wind belt. Radial increases in specificgravity of the type documented here may be important in evaluatingthe carbon present in forest stores. Copyright 2000 Annals ofBotany Company Buttresses, wood specific gravity, vessel diameter and density, Elaeocarpus angustifolius.     

Wood Specific Gravity Variations and Biomass of Central African Tree Species: The Simple Choice of the Outer Wood

Context

Wood specific gravity is a key element in tropical forest ecology. It integrates many aspects of tree mechanical properties and functioning and is an important predictor of tree biomass. Wood specific gravity varies widely among and within species and also within individual trees. Notably, contrasted patterns of radial variation of wood specific gravity have been demonstrated and related to regeneration guilds (light demanding vs. shade-bearing). However, although being repeatedly invoked as a potential source of error when estimating the biomass of trees, both intraspecific and radial variations remain little studied. In this study we characterized detailed pith-to-bark wood specific gravity profiles among contrasted species prominently contributing to the biomass of the forest, i.e., the dominant species, and we quantified the consequences of such variations on the biomass.

Methods

Radial profiles of wood density at 8% moisture content were compiled for 14 dominant species in the Democratic Republic of Congo, adapting a unique 3D X-ray scanning technique at very high spatial resolution on core samples. Mean wood density estimates were validated by water displacement measurements. Wood density profiles were converted to wood specific gravity and linear mixed models were used to decompose the radial variance. Potential errors in biomass estimation were assessed by comparing the biomass estimated from the wood specific gravity measured from pith-to-bark profiles, from global repositories, and from partial information (outer wood or inner wood).

Results

Wood specific gravity profiles from pith-to-bark presented positive, neutral and negative trends. Positive trends mainly characterized light-demanding species, increasing up to 1.8 g.cm^-3 per meter for Piptadeniastrum africanum, and negative trends characterized shade-bearing species, decreasing up to 1 g.cm^-3 per meter for Strombosia pustulata. The linear mixed model showed the greater part of wood specific gravity variance was explained by species only (45%) followed by a redundant part between species and regeneration guilds (36%). Despite substantial variation in wood specific gravity profiles among species and regeneration guilds, we found that values from the outer wood were strongly correlated to values from the whole profile, without any significant bias. In addition, we found that wood specific gravity from the DRYAD global repository may strongly differ depending on the species (up to 40% for Dialium pachyphyllum).

Main Conclusion

Therefore, when estimating forest biomass in specific sites, we recommend the systematic collection of outer wood samples on dominant species. This should prevent the main errors in biomass estimations resulting from wood specific gravity and allow for the collection of new information to explore the intraspecific variation of mechanical properties of trees.     

The facultative bimodal growth pattern in Quercus ilex – A simple model to predict sub-seasonal and inter-annual growth

In Mediterranean climates, bimodal growth patterns, corresponding to two peaks in radial increment during favorable seasons, have been described in several tree species. However, we lack a better mechanistic understanding of bimodality and its potential responses to the predicted warming and aridification trends. Filling this research gap is important since growth duration affects the capacity of trees to form wood and uptake carbon. Here we used an 11-year (1994–2004) long record of dendrometer data of the Mediterranean Holm oak (Quercus ilex) and compared how climate related to radial increment in trees from the south- and the north-facing slopes. We also related climate variables to tree-ring width and the production of intra-annual density fluctuations (IADFs), which reflects bimodality. In this paper, we introduce a model called VS-Lite2 to simulate tree-growth dynamics, which is a modified version of the process-based Vaganov-Shashkin Lite model. The VS-Lite2 model adequately reproduced the bimodal intra‐annual pattern of radial growth, IADFs, and annual tree growth. Trees from the south-oriented slope grew more, produced more IADFs and showed a more marked bimodal pattern than trees from the north-facing slope. These differences agree with the observation that late-summer drought constrained growth. Therefore, radial-growth models should consider plastic bimodality and micro-environmental conditions in areas subjected to seasonal droughts.     

Wood specific gravity and anatomy in Heliocarpus appendiculatus (Tiliaceae)

Specific gravity exhibits extremely large radial increases with distance from the pith in Heliocarpus appendiculatus Turcz. (Tiliaceae), a pioneer of neotropical wet forests. To determine some of the wood anatomical changes associated with this increase, wood samples taken at breast height from three trees were divided into 1.0-cm-long segments from pith to bark. Measurements were made of fiber wall thickness, fiber lumen diameter, and percentages of fibers, axial parenchyma, ray parenchyma, and vessels on sections prepared from each segment. The extreme radial increases in specific gravity were associated with increases in fiber wall thickness, decreases in fiber diameter, decreases in fiber lumen diameter, and changes in the relative proportions of fibers and parenchyma. The increase in percent fiber concomitant with a decrease in axial parenchyma was the most important contributor to the increase in specific gravity in this species. The best predictor of specific gravity was percent fibers (r = 0.91, 0.92, 0.94) or percent axial parenchyma (r = -0.92, -0.91, -0.95), two variables that were highly intercorrelated (r = -0.95, -0.98, -0.99).     

Effects of bending stress on taper and growth of stems of youngEucalyptus regnans trees

A glasshouse experiment investigated the effect of bending stress on stem radial and height growth and stem taper ofEucalyptus regnans seedlings. Eighteen-week-old, potted seedlings were bent continuously for 8 weeks with a static bending stress. The bending treatment was then removed and the seedlings grown for another 12 weeks. Other seedlings were stayed vertically throughout the experiment whilst control seedlings were neither bent nor stayed. Seedlings were rotated every 2 days to prevent reaction wood developing asymmetrically in the stems of bent trees. Bent trees had higher radial growth rates, developed more tapered stems and had higher safety factors (the ratio of stem radius to the minimum radius required to prevent the tree toppling over) than unbent seedlings. They produced a band of tension wood in their stems and ceased height growth whilst bent. When bending ceased, they resumed normal radial and height growth. Unbent trees developed more cylindrical stems. There were no differences in growth behaviour between stayed and control trees. Bent and unbent trees all developed a butt swell, the taper of which was not affected by treatment. It was concluded that bending stress has substantial effects on both the size and taper of tree stems. However, the development of butt swell is independent of the bending stress applied. The results were considered in relation to biomechanical theories of tree stem development.     

Architecture and size relations: an essay on the apple (Malus x domestica, Rosaceae) tree

The influence of tree size independent of age on some architectural features (annual shoot length, lateral branching, flowering) was investigated on 4-yr-old apple (Malus × domestica) trees either own-rooted or grafted on the dwarfing rootstock M.9, giving rise to large and small trees, respectively. Tree size significantly affected the length of the first annual shoot of bottom branches with a lesser effect on the subsequent annual shoots of the same branches and on branches situated higher in the tree canopy. The linear regression parameters, i.e., slopes and intercepts, between annual shoot length and number of growing laterals were affected by the genotype and, depending on genotype, by tree size. Flowering was generally lower, delayed, and more irregular on large trees compared to small trees, with on average similar ranking of genotypes regardless of tree size. This study provides evidence for a specific effect of tree size, as affected by the root system, on architectural development of the apple tree regardless of the genotype. From an architectural viewpoint, the dwarfing mechanism could be interpreted as a faster physiological aging essentially related to the reduction in length of the first annual shoot of bottom branches and the high flowering on this shoot.     

Replicated throughfall exclusion experiment in an Indonesian perhumid rainforest: wood production,litter fall and fine root growth under simulated drought

Climate change scenarios predict increases in the frequency and duration of ENSO‐related droughts for parts of South‐East Asia until the end of this century exposing the remaining rainforests to increasing drought risk. A pan‐tropical review of recorded drought‐related tree mortalities in more than 100 monitoring plots before, during and after drought events suggested a higher drought‐vulnerability of trees in South‐East Asian than in Amazonian forests. Here, we present the results of a replicated (n = 3 plots) throughfall exclusion experiment in a perhumid tropical rainforest in Sulawesi, Indonesia. In this first large‐scale roof experiment outside semihumid eastern Amazonia, 60% of the throughfall was displaced during the first 8 months and 80% during the subsequent 17 months, exposing the forest to severe soil desiccation for about 17 months. In the experiment's second year, wood production decreased on average by 40% with largely different responses of the tree families (ranging from −100 to +100% change). Most sensitive were trees with high radial growth rates under moist conditions. In contrast, tree height was only a secondary factor and wood specific gravity had no influence on growth sensitivity. Fine root biomass was reduced by 35% after 25 months of soil desiccation while fine root necromass increased by 250% indicating elevated fine root mortality. Cumulative aboveground litter production was not significantly reduced in this period. The trees from this Indonesian perhumid rainforest revealed similar responses of wood and litter production and root dynamics as those in two semihumid Amazonian forests subjected to experimental drought. We conclude that trees from paleo‐ or neotropical forests growing in semihumid or perhumid climates may not differ systematically in their growth sensitivity and vitality under sublethal drought stress. Drought vulnerability may depend more on stem cambial activity in moist periods than on tree height or wood specific gravity.     

不同坡向川西亚高山林木竞争与叶片表型可塑性的关系研究

表型可塑性是植物生长响应外界环境变化的重要表现形式,体现了植物个体在环境胁迫下的适合度。但是关于植物表型可塑性的驱动机制仍然存在很多争议。为了探讨植物表型可塑性的影响因素,以四川省阿坝藏族羌族自治州位于同一海拔梯度但坡向相反的天然次生林为研究对象,分析了不同坡向竞争强度与10种树木叶片功能性状表型可塑性的关系的差异。研究发现：(1)研究样地中阴坡水分和养分资源优于阳坡;(2)阴坡上林木平均种内和种间竞争强度高于阳坡,阴坡上林木种内竞争强度随着树木个体大小的增加而显著性减少,阳坡上林木种内竞争强度却随着个体大小的增加而增加;(3)阴坡上叶片表型可塑性高于阳坡,表型可塑性随着个体大小的增加而增加,在阳坡上却随着个体大小增加而降低。这些结果表明阴坡上水分等资源环境条件优于阳坡,林木生长受到环境资源限制较少。在林木生长过程中,较高的竞争强度引起的资源重叠加剧,尤其是种内竞争强度的变化,从而导致了阴坡上较高的叶片表型可塑性。因为较高的竞争强度,随着林木个体大小的增加,树木需要更高的可塑性赢得竞争优势来获取更多的资源支持生长。但是在阳坡上,资源相对缺乏,环境资源对树木生长的限制降低了叶片表型的可塑...     

Comparative Studies on Woods of Three Species of Normal and Dwarf Trees

Wood structure of three species i.e. Larix chinensis Beissn., Alnus sibirica Fisch. and Quercus mongolica Fisch. of normal and dwarf trees were comparatively studied. The age variation of width of annual rings and size of tracheary elements in these trees follows their own regularities respectively during the course of ontogenesis. In normal, as trees grow, the width of annual rings increases gradually, reaches maximum, then declines in subsequence; while the size of tracheary elements exhibits a crescendo increment, and remains constant after reaching maximum. These trends were not affected by the change of enviromental conditions in which the trees grow. However, the width of annual rings and size of tracheary elements of dwarf trees which grow in adverse circumstances are respectively narrower and smaller than those of normal trees. Further, in dwarf tree of L. chinensis (coniferous wood); the diameter of bordered pits on tracheids of early wood decreases, ray height is lower and ray frequence is higher. In dwarf tree of A. sibirica (diffuse porous wood), vessel frequence increases and the number of bars per perforation decreases. The quatitative features of wood structure of dwarf trees, as compared with those of normal trees, showed more variations; whereas, their own qualitative features of wood structure remain constant. These findings might indicate that modifications of wood structure responsing directly to change of enviromental conditions are usually only qualitative.     

Three-dimensional distribution of mass movement signals in disturbed Picea abies (L.) Karst.

Dendrogeomorphologic approaches enable annual absolute dating of various potentially hazardous geomorphic processes. This study tested the dating precision of three commonly used dendrogeomorphic approaches used for the extraction of geomorphic signals from the tree ring series of disturbed trees. The first approach is based on the onset of reaction wood occurrence, and the other two approaches are based on the detection of abrupt changes in tree ring eccentricity. Nine 165 cm long stems of maximally 27 year old Norway spruces (Picea abies (L.) Karst.) growing on unstable slope affected by mass movement events of known age were analysed. Several sampling directions on stems were analysed to reveal the best sampling strategy for each tested approach. The relationships between stem curvature and the dating precision of the approaches were also analysed. The results of dating precision showed crucial differences between the tested approaches. Tree stem curvature did not influence dating precision. Overall, we concluded that the approaches based on eccentric increments are less precise than those based on reaction wood analysis when juvenile P. abies trees that grow on landslide slope are sampled. The importance of this study is in providing new insight into dating precision by analysing the whole tree stems of trees influenced by mass movements. This is unique since the approaches based on eccentricity increments have not been previously tested in such a detail.     

Modelling of auxin transport affected by gravity and differential radial growth

When a tree stem deviates from verticality, as a result of different environmental factors, patterns of differential radial growth appear. Higher rates of wood production have been observed on the lower side of the tree and lower rates in the opposite side. Biological studies on plant hormones have shown that the concentration of auxin induces radial growth. They also have demonstrated the redistribution of auxin transport in response to gravity. Auxin is then designated as a mediator for differential growth. This paper presents a model for three-dimensional (3-D) auxin transport in conifer trees, which includes gravity dependence. We obtain realistic heterogeneous patterns of auxin distribution over the tree. Then, we propose a law of growth based on auxin concentration to simulate successive differential radial growths. The predicted growths are compared with experimental results of reconstruction of 3-D annual growth of Radiata pine.     

三种正常与矮化植物的木材比较观察

本文比较观察了三种不同木材类型的正常与矮化树木的木材结构:针叶材的太白红杉(Larix chinensis Beissn.)、散孔材的辽东桤木(Alnus sibirica Fisch.)和环孔材的蒙古栎(Quercus mongolica Fisch.)。这些植物在个体发育过程中,各年的年轮宽度及管状分子长度、宽度等变化,存在各自的规律性。这些规律并不因生长环境的差异而变更。不过,逆境下生长的矮化植株的年轮宽度和管状分子的大小明显地比正常的要窄小,同时在木材结构的其它数量性状上,也有较大变化;然而,其各自的种属定性特征仍保持稳定。这些似乎可说明:生态环境的变化对木材结构的数量性状有较大影响,但是对木材种属的定性特征则无明显作用。     

Radial Gradients in Wood Specific Gravity,Water and Gas Content in Trees of a Mexican Tropical Rain Forest

Cell walls, water, and gas that have mechanical and physiological functions in wood, and wood specific gravity (WSG) is related to demographic traits. To understand variation in wood structure and function, we analyzed radial changes in WSG, in the gas and the water fractions from trees growing in four different habitats in a southern Mexican rain forest. Mean WSG was 0.55 ± 0.16, slightly lower than reported for other tropical forests. In 27 species, WSG decreased and in two species, it increased from pith to bark with a strong (r² = 0.65) negative correlation between WSG in the center of the tree and the radial WSG gradient. Habitat had some effect on mean WSG and trees growing on karst had significantly higher WSG than the same species growing on alluvial soil. The cell wall, water, and gas fractions accounted for 35 percent (range: 16–50), 42 percent (28–65), and 23 percent (2–56), respectively, of wood volume, with a negative correlation between the gas and the cell wall and between the gas and the water fractions, but not between the cell wall and water fractions. Radially increasing WSG is advantageous for pioneer trees with fast initial growth. We found that the water displacement method may result in biased WSG estimates. To increase the accuracy of WSG data, we suggest to measure sample volume geometrically using a constant diameter (that of the borer tip), to include radial variation in WSG, and to consider for possible site effects on species‐specific WSG.     

Vessel feature changes as a tool for detecting annual rings in tropical trees

Radial variations in vessel features (vessel lumen area and frequency) of four tropical tree species grown in tropical savanna, monsoon, and rainforest climates were investigated to detect indistinct annual rings. Leaf and soil water potentials were measured periodically to show annual variations in water availability and their relationship to radial variations in vessel features. In addition, the accuracy of annual-ring detection was estimated using trees of known age. Radial variations in vessel features showed annual cyclicity in all three sites. The vessel feature that showed clear annual cyclicity was different among the species and sites. Furthermore, the variation pattern of vessel features in the rainforest site tended to be different between two radii within individual trees, implying low synchronicity of wood formation in the wet area. Compared with leaf and soil water potentials, vessel features varied independent of water availability in all species in the rainforest site and most species in the savanna and monsoon sites. The direct effect of low water availability on vessel features was considered only in one species in the savanna and monsoon sites. Nevertheless, the deviation of vessel feature cycles from tree age was similar and reasonably small (mostly within ±10 %) among all sites. These results indicated a circannual rhythm in vessel formation. Thus, vessel feature changes seemed to aid in detecting annual rings in trees throughout wide tropical areas; however, asynchronous wood formation must be considered in dendrochronology, especially in humid tropics.     

关帝山云杉次生林树木径向生长的空间关联及其影响因子

以关帝山4 hm²云杉次生林样地为研究对象,按照CTFS(Center for Tropical Forest Science)技术规范对样地树木进行连续定位监测。利用2010至2015年间样地主要树种生长量观测数据,结合地形、土壤等环境因子调查及采样测定数据,分析了树木种群径向生长的空间关联性及其随生境的变化,并探讨了树木种群径向生长的影响因素。结果表明,青杄、华北落叶松、红桦、白桦和辽东栎为云杉次生林主要树种,在样地4个生境型(山脊生境、低海拔缓坡生境、高海拔缓坡生境、低洼地陡坡生境)中均有分布且呈现不同的径级结构。标记相关函数分析显示,同一生境型中,5树种径向生长的空间关联性各异;对于同一树种,径向生长的空间自相关性不仅具有尺度依赖性,同时生境型的不同导致树木径向生长的空间关联性发生变化。线性混合效应模型分析显示,初始胸径对树木径向生长的显著正效应在样地各类生境型的所有种群中普遍存在;生物因子对树木径向生长的显著影响只在特定生境型的青杄种群中被检测到,表明树木径向生长受同种邻体影响,但其影响显著性因树种而异;环境因子中,海拔和凹凸度对树木径向生长呈显著负效应...