The links between wood traits and species demography change during tree development in a lowland tropical rainforest.

One foundational assumption of trait-based ecology is that traits can predict species demography. However, the links between traits and demographic rates are, in general, not as strong as expected. These weak associations may be due to the use of traits that are distantly related to performance, and/or the lack of consideration of size-related variations in both traits and demographic rates. Here, we examined how wood traits were related to demographic rates in 19 tree species from a lowland forest in eastern Amazonia. We measured 11 wood traits (i.e. structural, anatomical and chemical traits) in sapling, juvenile and adult wood; and related them to growth and mortality rates (MR) at different ontogenetic stages. The links between wood traits and demographic rates changed during tree development. At the sapling stage, relative growth rates (RGR) were negatively related to wood specific gravity (WSG) and total parenchyma fractions, while MR decreased with radial parenchyma fractions, but increased with vessel lumen area (VA). Juvenile RGR were unrelated to wood traits, whereas juvenile MR were negatively related to WSG and axial parenchyma fractions. At the adult stage, RGR scaled with VA and wood potassium concentrations. Adult MR were not predicted by any trait. Overall, the strength of the trait-demography associations decreased at later ontogenetic stages. Our results indicate that the associations between traits and demographic rates can change as trees age. Also, wood chemical or anatomical traits may be better predictors of growth and MR than WSG. Our findings are important to expand our knowledge on tree life-history variations and community dynamics in tropical forests, by broadening our understanding on the links between wood traits and demography during tree development.

Earlier onset and slower heartwood investment in faster-growing trees of African tropical species.

BACKGROUND AND AIMS: Heartwood plays an important role in maintaining the structural integrity of trees. While, its formation has long be thought to be solely driven by internal ageing processes, more recent hypotheses suggest that heartwood formation acts as a regulator of the tree water balance by modulating sapwood quantities. Testing both hypotheses would shed light on the potential ecophysiological nature of heartwood formation, a very common process in trees. METHODS: We measured heartwood and sapwood quantities, xylem conduits and growth ring width and number on 406 stems of Pericopsis elata with ages ranging from 2 to 237 years. A subset of 17 trees with similar ages but varying growth rate were sampled in a shaded (slower growth) and sun-exposed (faster growth) site. We used regression analysis and structural equations modelling to investigate the dynamics and drivers of heartwood formation. KEY RESULTS: We found a positive effect of growth rate on the probability of heartwood occurrence, suggesting an earlier heartwood onset in faster-growing stems. After this onset age, heartwood area increases with stem diameter and age. Despite the similar heartwood production per unit stem diameter increment, shaded trees produce heartwood faster than sun-exposed trees. Tree age and hydraulics showed similar direct effects on heartwood and sapwood area of sun-exposed trees, suggesting their mutual role in driving the heartwood dynamics of sun-exposed trees. However, for shaded trees, only tree hydraulics showed a direct effect, suggesting its prominent role over age in driving the heartwood dynamics in limited growing conditions. The positive relationship between growth rate and maximum stomatal conductance supports this conclusion. CONCLUSIONS: Heartwood area increases as the tree ages but at a slower rate in trees where water demand is balanced by a sufficient water supply. Our findings suggest that heartwood formation is not only a structural but also a functional process.

Microspines in tropical climbing plants: a small-scale fix for life in an obstacle course.

Many climbing plants have microspines on their stems, which facilitate attachment and prevent slipping and falling from host plant supports. Extending via growth through complex environments and anchoring stems to substrates with minimal contact forces are key benefits for climbing plants. Microspines are also highly desirable features for new technologies and applications in soft robotics. Using a novel sled-like device, we investigated static and sliding attachment forces generated by stems in 10 species of tropical climber from French Guiana differing in size and climbing habit. Eight species showed higher static and sliding forces when their stems were pulled in the basal direction against a standard surface than in the apical direction. This anisotropic behaviour suggests that tropical climbers have evolved different ratchet-like mechanisms that allow easy sliding forwards but are resistant to slipping downwards. The presence of a downwards 'stick-and-slip' phenomenon, where static attachment is not significantly stronger than maximal sliding attachment, was present in most species apart from three showing relatively weak attachment by microspines. This indicates that diverse microspine attachment strategies exist in climbing plants. This diversity of functional properties offers a range of potential design specifications for climbing strategies on different substrates for artificial climbing artefacts.

High-Resolution X-Ray Computed Tomography: A New Workflow for the Analysis of Xylogenesis and Intra-Seasonal Wood Biomass Production.

Understanding tree growth and carbon sequestration are of crucial interest to forecast the feedback of forests to climate change. To have a global understanding of the wood formation, it is necessary to develop new methodologies for xylogenesis measurements, valid across diverse wood structures and applicable to both angiosperms and gymnosperms. In this study, the authors present a new workflow to study xylogenesis using high-resolution X-ray computed tomography (HRXCT), which is generic and offers high potential for automatization. The HXRCT-based approach was benchmarked with the current classical approach (microtomy) on three tree species with contrasted wood anatomy (Pinus nigra, Fagus sylvatica, and Quercus robur). HRXCT proved to estimate the relevant xylogenesis parameters (timing, duration, and growth rates) across species with high accuracy. HRXCT showed to be an efficient avenue to investigate tree xylogenesis for a wide range of wood anatomies, structures, and species. HRXCT also showed its potential to provide quantification of intra-annual dynamics of biomass production through high-resolution 3D mapping of wood biomass within the forming growth ring.

How does bark contribution to postural control change during tree ontogeny? A study of six Amazonian tree species.

Recent works revealed that bark is able to produce mechanical stress to control the orientation of young tilted stems. Here we report how the potential performance of this function changes with stem size in six Amazonian species with contrasted bark anatomy. The potential performance of the mechanism depends both on the magnitude of bark stress and the relative thickness of the bark. We measured bark longitudinal residual strain and density, and the allometric relationship between bark thickness and stem radius over a gradient of tree sizes. Constant tensile stress was found in species that rely on bark for the control of stem orientation in young stages. Other species had increasing compressive stress, associated with increasing density attributed to the development of sclereids. Compressive stress was also associated with low relative bark thickness. The relative thickness of bark decreased with size in all species, suggesting that a reorientation mechanism based on bark progressively performs less well as the tree grows. However, greater relative thickness was observed in species with more tensile stress, thereby evidencing that this reduction in performance is mitigated in species that rely on bark for reorientation.

Leveraging Signatures of Plant Functional Strategies in Wood Density Profiles of African Trees to Correct Mass Estimations From Terrestrial Laser Data.

Wood density (WD) relates to important tree functions such as stem mechanics and resistance against pathogens. This functional trait can exhibit high intraindividual variability both radially and vertically. With the rise of LiDAR-based methodologies allowing nondestructive tree volume estimations, failing to account for WD variations related to tree function and biomass investment strategies may lead to large systematic bias in AGB estimations. Here, we use a unique destructive dataset from 822 trees belonging to 51 phylogenetically dispersed tree species harvested across forest types in Central Africa to determine vertical gradients in WD from the stump to the branch tips, how these gradients relate to regeneration guilds and their implications for AGB estimations. We find that decreasing WD from the tree base to the branch tips is characteristic of shade-tolerant species, while light-demanding and pioneer species exhibit stationary or increasing vertical trends. Across all species, the WD range is narrower in tree crowns than at the tree base, reflecting more similar physiological and mechanical constraints in the canopy. Vertical gradients in WD induce significant bias (10%) in AGB estimates when using database-derived species-average WD data. However, the correlation between the vertical gradients and basal WD allows the derivation of general correction models. With the ongoing development of remote sensing products providing 3D information for entire trees and forest stands, our findings indicate promising ways to improve greenhouse gas accounting in tropical countries and advance our understanding of adaptive strategies allowing trees to grow and survive in dense rainforests.

Mechanical contribution of secondary phloem to postural control in trees: the bark side of the force.

To grow straight, plants need a motor system that controls posture by generating forces to offset gravity. This motor function in trees was long thought to be only controlled by internal forces induced in wood. Here we provide evidence that bark is involved in the generation of mechanical stresses in several tree species. Saplings of nine tropical species were grown tilted and staked in a shadehouse and the change in curvature of the stem was measured after releasing from the pole and after removing the bark. This first experiment evidenced the contribution of bark in the up-righting movement of tree stems. Combined mechanical measurements of released strains on adult trees and microstructural observations in both transverse and longitudinal/tangential plane enabled us to identify the mechanism responsible for the development of asymmetric mechanical stress in the bark of stems of these species. This mechanism does not result from cell wall maturation like in wood, or from the direct action of turgor pressure like in unlignified organs, but is the consequence of the interaction between wood radial pressure and a smartly organized trellis structure in the inner bark.

The pipe model theory half a century on: a review.

Background: More than a half century ago, Shinozaki et al. (Shinozaki K, Yoda K, Hozumi K, Kira T. 1964a. A quantitative analysis of plant form - the pipe model theory. I. Basic analyses. Japanese Journal of Ecology B: 97-105) proposed an elegant conceptual framework, the pipe model theory (PMT), to interpret the observed linear relationship between the amount of stem tissue and corresponding supported leaves. The PMT brought a satisfactory answer to two vividly debated problems that were unresolved at the moment of its publication: (1) What determines tree form and which rules drive biomass allocation to the foliar versus stem compartments in plants? (2) How can foliar area or mass in an individual plant, in a stand or at even larger scales be estimated? Since its initial formulation, the PMT has been reinterpreted and used in applications, and has undoubtedly become an important milestone in the mathematical interpretation of plant form and functioning. Scope: This article aims to review the PMT by going back to its initial formulation, stating its explicit and implicit properties and discussing them in the light of current biological knowledge and experimental evidence in order to identify the validity and range of applicability of the theory. We also discuss the use of the theory in tree biomechanics and hydraulics as well as in functional-structural plant modelling. Conclusions: Scrutinizing the PMT in the light of modern biological knowledge revealed that most of its properties are not valid as a general rule. The hydraulic framework derived from the PMT has attracted much more attention than its mechanical counterpart and implies that only the conductive portion of a stem cross-section should be proportional to the supported foliage amount rather than the whole of it. The facts that this conductive portion is experimentally difficult to measure and varies with environmental conditions and tree ontogeny might cause the commonly reported non-linear relationships between foliage and stem metrics. Nevertheless, the PMT can still be considered as a portfolio of properties providing a unified framework to integrate and analyse functional-structural relationships.