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.     

Wood density and vessel traits as distinct correlates of ecological strategy in 51 California coast range angiosperms

Wood density and vessel characteristics are functionally interrelated, yet they may have distinct ecological associations. In a comparative study of 51 angiosperm species ranging from chaparral shrubs to riparian trees, we examined relationships among wood density and vessel traits and their ecological correlates. Mean vessel lumen area and vessel density (number mm(-2)) varied widely (7- to 10-fold). In multivariate analyses, both vessel traits were negatively correlated with wood density, which varied more narrowly (< 2-fold). Vessel density and lumen area were inversely related across species, allowing a broad range of vessel traits within a narrow range of wood density. Phylogenetic independent contrasts indicated correlated inverse evolutionary change in vessel traits. Each trait had a distinct pattern of ecological correlation -- wood density was most strongly associated with soil water, and vessel traits showed contrasting relationships with plant height. Within a narrow range of wood density, there was significant variation in vessel traits. Given their particular ecological associations, the results suggest that wood density and vessel traits describe two distinct ecological axes.     

Wood mechanics, allometry, and life-history variation in a tropical rain forest tree community

Wood density plays a central role in the life-history variation of trees, and has important consequences for mechanical properties of wood, stem and branches, and tree architecture. Wood density, modulus of rupture, modulus of elasticity, and safety factors for buckling and bending were determined for saplings of 30 Bolivian rain forest tree species, and related to two important life-history axes: juvenile light demand and maximum adult stature. Wood density was strongly positively related to wood strength and stiffness. Species safety factor for buckling was positively related to wood density and stiffness, but tree architecture (height : diameter ratio) was the strongest determinant of mechanical safety. Shade-tolerant species had dense and tough wood to enhance survival in the understorey, whereas pioneer species had low-density wood and low safety margins to enhance growth in gaps. Pioneer and shade-tolerant species showed opposite relationships between species traits and adult stature. Light demand and adult stature affect wood properties, tree architecture and plant performance in different ways, contributing to the coexistence of rain forest species.     

Wood anatomy and wood density in shrubs: Responses to varying aridity along transcontinental transects

Wood density plays a key role in ecological strategies and life history variation in woody plants, but little is known about its anatomical basis in shrubs. We quantified the relationships between wood density, anatomy, and climate in 61 shrub species from eight field sites along latitudinal belts between 31° and 35° in North and South America. Measurements included cell dimensions, transverse areas of each xylem cell type and percentage contact between different cell types and vessels. Wood density was more significantly correlated with precipitation and aridity than with temperature. High wood density was achieved through reductions in cell size and increases in the proportion of wall relative to lumen. Wood density was independent of vessel traits, suggesting that this trait does not impose conduction limitations in shrubs. The proportion of fibers in direct contact with vessels decreased with and was independent of wood density, indicating that the number of fiber-vessel contacts does not explain the previously observed correlation between wood density and implosion resistance. Axial and radial parenchyma each had a significant but opposite association with wood density. Fiber size and wall thickness link wood density, life history, and ecological strategies by controlling the proportion of carbon invested per unit stem volume.     

In the lack of extreme pioneers: trait relationships and ecological strategies of 66 subtropical tree species

Aims Despite the growing interest in the topic of functional ecology, there are still forest regions that have not been examined, as most work has been done in the tropics. Unresolved issues include the strength of a growth-mortality trade-off in trees (originally identified for seedlings) and the nature of the association between plant traits and vital rates, if any. Our objectives were to examine whether (i) ecological strategies in South American mixed forests are organized along the fast competitor × slow stress-tolerator and height gradients as the main strategy axes depicted in the overall trait and vital rate correlation structure, and (ii) a tentative path model we proposed can explain the patterns of covariation among traits and vital rates.Methods We studied a different habitat (subtropical mixed conifer-hardwood forests) and region (Brazilian Atlantic Forest) from the majority of related studies in forests, carried out in the Neotropical region. Data on total height, stem slenderness, crown depth, wood density, specific leaf area, leaf and seed length, seed dispersal mode, annual mortality, diameter relative growth rate and relative growth rate under favorable conditions were measured in southern Brazil for 66 tree species. Data were subjected to principal components analysis and path analysis. Restricted data on saplings and treelets were analyzed through correlation.Important findings Studied traits were reduced to four principal components. Principal components analysis produced axes that fit the resource acquisition versus resource-conservation and the height-mortality trade-offs, although the former was split into two distinct axes. Seed size and seed dispersal mode appeared independently of these axes. A path model showed that leaf length and specific leaf area caused direct changes in trunk slenderness and, indirectly through growth, affected mortality. Expected trade-offs between growth and survivorship and between wood density and stem slenderness trade-offs were not found. This may result from the lack of extreme pioneers and over-representation of slow-growing hardwood species found in Atlantic subtropical forests of South America. This suggests that the fastest growing species in the region do not grow so fast as to compromise wood density and survivorship, but grow fast enough to benefit from increased size. Relationships between traits and vital rates seem to be mediated by the assembly process of regional floras, and the relative importance of traits like SLA and wood density may vary between floristic regions.     

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.     

SURFACE AREA RELATIONS OF WOODY PLANTS AND FOREST COMMUNITIES

Surface area of wood and bark is an important dimension of forests, with implications for respiration rate, energy exchange, and water and mineral budgets. Surface area of stem wood and bark can be estimated effectively from linear regressions on conic surface (one-half basal circumference times tree height) or from regressions of the logarithm of area on the logarithm of diameter at breast height. Branch surface can be estimated from a formula using branch basal diameter, length, and number of current twigs, and from logarithmic regressions of branch bark surface on basal diameter of branches and breast-height diameter of trees. In temperate deciduous forests several square meters of plant surface occur above each square meter of ground surface; these plant surfaces include 0.3–0.6 m² of stem bark, 1.2–2.2 m² of branch bark, and 3.0–6.0 m² of leaf blades. Branch bark surface increases more rapidly than leaf surface with increasing size of branches and trees. Growth and aging of trees, and maturation of forests, imply increasing ratios of bark (and wood) surface to the photosynthetic leaf surface which supports its growth and respiration.     

Ecological and evolutionary determinants of a key plant functional trait: wood density and its community-wide variation across latitude and elevation

Wood density is an important trait in trees indicative of their life history and mechanical and physiological strategies. The following examines the evolutionary ecology of wood density using a large database for seed plants. In particular, we focused on the geographic and phylogenetic variation in wood density for both gymnosperms and angiosperms. A phylogenetic supertree was constructed for over 4600 taxa, allowing for comprehensive analyses of divergences across the seed plant phylogeny. Community-wide means and variances for wood densities were quantified for 171 standardized forest communities. Wood density was generally highly conserved across the entire seed plant phylogeny, yet large divergences were found within the rosid clade. Geographic and community variation in wood density, however, was significantly lower in temperate and high elevation communities, dominated by gymnosperms, than in tropical lowland communities, dominated by angiosperms, suggesting an increase in trait and, to some extent, clade filtering with latitude and elevation. Together, our results support the notion that both biotic and abiotic forces have been important in the evolution of wood density as well as in controlling the observed trait mean and variance across geographic gradients.     

Molecular Genetics of Growth and Development in Populus. IV. Mapping Qtls with Large Effects on Growth, Form, and Phenology Traits in a Forest Tree

We have mapped quantitative trait loci (QTLs) for commercially important traits (stem growth and form) and an adaptive trait (spring leaf flush) in a Populus F(2) generation derived from a cross between interspecific F(1) hybrids (P. trichocarpa X P. deltoides). Phenotypic data were collected over a 2-year period from a replicated clonal trial containing ramets of the parental, F(1), and F(2) trees. Contrary to the assumptions of simple polygenic models of quantitative trait inheritance, 1-5 QTLs of large effect are responsible for a large portion of the genetic variance in each of the traits measured. For example, 44.7% of the genetic variance in stem volume after 2 years of growth is controlled by just two QTLs. QTLs governing stem basal area were found clustered with QTLs for sylleptic branch leaf area, sharing similar chromosomal position and mode of action and suggesting a pleiotropic effect of QTLs ultimately responsible for stem diameter growth.     

Roles of climate and functional traits in controlling toothed vs. untoothed leaf margins

? Premise of the study: Leaf-margin state (toothed vs. untoothed) forms the basis of several popular methods for reconstructing temperature. Some potential confounding factors have not been investigated with large data sets, limiting our understanding of the adaptive significance of leaf teeth and their reliability to reconstruct paleoclimate. Here we test the strength of correlations between leaf-margin state and deciduousness, leaf thickness, wood type (ring-porous vs. diffuse-porous), height within community, and several leaf economic variables. ? Methods: We assembled a trait database for 3549 species from six continents based on published and original data. The strength of associations between traits was quantified using correlational and principal axes approaches. ? Key results: Toothed species, independent of temperature, are more likely to be deciduous and to have thin leaves, a high leaf nitrogen concentration, a low leaf mass per area, and ring-porous wood. Canopy trees display the highest sensitivity between leaf-margin state and temperature; subcanopy plants, especially herbs, are less sensitive. ? Conclusions: Our data support hypotheses linking the adaptive significance of teeth to leaf thickness and deciduousness (in addition to temperature). Toothed species associate with the "fast-return" end of the leaf economic spectrum, providing another functional link to thin leaves and the deciduous habit. Accounting for these confounding factors should improve climate estimates from tooth-based methods.     

Spatial patterns of wood traits in China are controlled by phylogeny and the environment

Aim Wood properties are related to tree physiology and mechanical stability and are influenced by both phylogeny and the environment. However, it remains unclear to what extent geographical gradients in wood traits are shaped by either phylogeny or the environment. Here we aimed to disentangle the influences of phylogeny and the environment on spatial trends in wood traits. Location China. Methods We compiled a data set of 11 wood properties for 618 tree species from 98 sampling sites in China to assess their phylogenetic and spatial patterns, and to determine how many of the spatial patterns in wood properties are attributable to the environment after correction for phylogenetic influences. Result All wood traits examined exhibited significant phylogenetic signal. The widest divergence in wood traits was observed between gymnosperms and angiosperms, Rosids and Asterids, Magnoiliids and Eudicots, and in Lamiales. For most wood traits, the majority of trait variation was observed at genus and species levels. The mechanical properties of wood showed correlated evolution with wood density. Most of the mechanical properties of wood exhibited significant latitudinal variation but limited or no altitudinal variation, and were positively correlated with mean annual precipitation based on both Pearson's correlation analysis and the phylogenetic comparative method. Correlations at family level between mean annual temperature and wood density, compression strength, cross‐section hardness, modulus of elasticity and volumetric shrinkage coefficient became significant after correction for phylogenetic influences. Main conclusions Phylogeny interacted with the environment in shaping the spatial patterns of wood traits of trees across China because most wood properties showed strong phylogenetic conservatism and thus affected environmental tolerances and distributions of tree species. Mean annual precipitation was a key environmental factor explaining the spatial patterns of wood traits. Our study provides valuable insights into the geographical patterns in productivity, distribution and ecological strategy of trees linking to wood traits.     

Transgenic poplars with reduced lignin show impaired xylem conductivity, growth efficiency and survival

We studied xylem anatomy and hydraulic architecture in 14 transgenic insertion events and a control line of hybrid poplar (Populus spp.) that varied in lignin content. Transgenic events had different levels of down-regulation of two genes encoding 4-coumarate:coenzyme A ligase (4CL). Two-year-old trees were characterized after growing either as free-standing trees in the field or as supported by stakes in a greenhouse. In free-standing trees, a 20 to 40% reduction in lignin content was associated with increased xylem vulnerability to embolism, shoot dieback and mortality. In staked trees, the decreased biomechanical demands on the xylem was associated with increases in the leaf area to sapwood area ratio and wood specific conductivity (k(s)), and with decreased leaf-specific conductivity (k(l)). These shifts in hydraulic architecture suggest that the bending stresses perceived during growth can affect traits important for xylem water transport. Severe 4CL-downregulation resulted in the patchy formation of discoloured, brown wood with irregular vessels in which water transport was strongly impeded. These severely 4CL-downregulated trees had significantly lower growth efficiency (biomass/leaf area). These results underscore the necessity of adequate lignification for mechanical support of the stem, water transport, tree growth and survival.     

Vertical gradients in leaf trait diversity in a New Zealand forest

Leaves come in a remarkable diversity of sizes and shapes. However, spatial patterns in leaf trait diversity are rarely investigated and poorly resolved. We used a hierarchical approach to evaluate vertical variability in leaf morphology (i.e., leaf trait diversity) in 16 common tree and shrub species inhabiting a New Zealand forest. Height-related heterogeneity in leaf area, specific leaf area, circularity and length to width ratio was analyzed at three scales: (1) among leaves within plants, (2) among plants within species and (3) among species within functional groups (i.e., trees vs. shrubs). Results were scale dependent. Among-leaf morphological diversity was unrelated to plant height. Among-individual morphological diversity increased with the average height of each species, indicating that taller plant species express a greater range of leaf traits than shorter species. Among-species morphological diversity was higher in shrubs than in trees. We hypothesize that scale-dependent patterns in leaf trait diversity result from scale-dependent adaptations to forest environmental conditions. As trees grow from the forest floor into the canopy, they are exposed to a range of environmental conditions, which may select for a range of leaf traits through ontogeny. Conversely, shrubs never reach the forest canopy and may instead be differentially adapted to suites of environmental conditions associated with different stages of forest recovery from tree-fall disturbances. Overall results indicate that vertical patterns in leaf trait diversity exist. However, their strength and directionality are strongly scale-dependent, suggesting that different processes govern leaf shape diversity at different levels of ecological organization.     

Correlating Stem Biomechanical Properties of Hawaiian Canopy Trees with Hurricane Wind Damage1

Hawaiian forests are subject to the effects of periodic hurricane conditions. Hurricane Iniki struck the island of Kauai, Hawaii on September 11, 1992 with winds exceeding 200 km/h and caused defoliation, felling of trees by snapping and uprooting, and standing tree mortality due excessive limb and leaf loss. The purpose of this study was to evaluate if measured wood mechanical characteristics could be correlated with stem failure of trees under windstorm conditions. A field survey indicated that post-hurricane stem condition (snapped, uprooted, or standing) differed among five common canopy species and was significantly correlated with stem apparent elastic modulus (relative flexibility). Species that tended to snap had significantly higher apparent elastic moduli than those that remained standing or were uprooted. Wood density and stem diameter were not significantly related to stem failure mode. Native trees had a higher percentage per species of standing individuals but also had increased uprooting. Nonnative tree species were more often snapped and fewer were standing after the hurricane. The higher incidence of stem failure for introduced canopy trees may increase the spread of alien understory species following wind disturbance events. These relationships provide a simple means to predict relative differences in stem failure due to high wind conditions and should be considered in planning reforestation efforts on the Hawaiian Islands.     

Stem diameter growth rates in a fire‐prone savanna correlate with photosynthetic rate and branch‐scale biomass allocation,but not specific leaf area

Plant growth rates strongly determine ecosystem productivity and are a central element of plant ecological strategies. For laboratory and glasshouse‐grown seedlings, specific leaf area (SLA; ratio of leaf area to mass) is a key driver of interspecific variation in growth rate (GR). Consequently, SLA is often assumed to drive GR variation in field‐grown adult plants. However, there is an increasing evidence that this is not the general case. This suggests that GR – SLA relationships (and perhaps those for other traits) may vary depending on the age or size of the plants being studied. Here we investigated GR – trait relationships and their size dependence among 17 woody species from an open‐canopy, fire‐prone savanna in northern Australia. We tested the predictions that SLA and stem diameter growth rate would be positively correlated in saplings but unrelated in adults while, in both age classes, faster‐GR species would have higher light‐saturated photosynthetic rate (A_sat), higher leaf nutrient concentrations, higher branch‐scale biomass allocation to leaf versus stem tissues and lower wood density (WD). SLA showed no relationship to stem diameter GR, even in saplings, and the same was true of leaf N and P concentrations, and WD. However, branch‐scale leaf:stem allocation was strongly related to GR in both age groups, as was A_sat. Together, these two traits accounted for up to 80% of interspecific variation in adult GR, and 41% of sapling GR. A_sat is rarely measured in field‐based GR studies, and this is the first report of branch‐scale leaf:stem allocation (analogous to a benefit:cost ratio) in relation to plant growth rate. Our results suggest that we may yet find general trait‐drivers of field growth rates, but SLA will not be one.     

Does grazing induce intraspecific trait variation in plants from a sub‐humid mountain ecosystem?

Livestock grazing represents an important human disturbance for vegetation worldwide. We analysed the intraspecific differences in mean trait values between different grazing regimes (ungrazed and grazed) and explored whether these differences are consistent across species in a sub‐humid mountain ecosystem in Central Argentina. We selected 14 species of eight different families, co‐occurring in both regimes and comprising herbaceous (grasses and forbs) and woody (shrubs and trees) plants. For each species and grazing regime we measured 12 traits related to plant size, carbon fixation and water use. We found that plants in the grazed regime had consistently smaller leaves and shorter stature and internodal length than plants of the same species under the ungrazed regime. For the remaining traits the responses were species‐specific. Dry matter content, leaf tensile strength and minimum leaf water potential (Ψ_leaf) showed contrasting responses to grazing. Specific leaf area, wood density and potential water content of wood showed almost no significant responses except for very few species. Neither leaf area per shoot mass nor leaf area per sapwood area differed significantly between grazing regimes. Our study suggested that the intraspecific variation found for the size‐related traits would allow species to respond to grazing without modifying markedly other structural traits, a plastic response that might increase the probability of species success.     

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.

     

Leaf traits and their interrelationship of 23 plant species in southeast of Keerqin Sandy Lands,China

Six leaf traits,i.e.,fresh mass (FM),dry mass (DM),leaf dry matter content (DMC),area (AR),specific leaf area (SLA) and thickness (TH) from 23 plant species in the southeastern Keerqin Sandy Lands,China were measured.The results show that leaf traits of herbs were more diversified than those of shrubs and trees and aver-age SLA tended towards a decreasing trend from herbs to shrubs to trees.On the contrary,DMC and DM show an upward trend from herbs to shrubs to trees.No apparent difference was found in TH.Except for DM and TH,there were significant variations in SLA and DMC among three different growth forms.Moreover,a significant correla-tion was found between SLA and DMC.It is concluded that SLA and DMC could be used to predict species posi-tion along a resource use gradient.