The contribution of insects to global forest deadwood decomposition.

The amount of carbon stored in deadwood is equivalent to about 8 per cent of the global forest carbon stocks1. The decomposition of deadwood is largely governed by climate2-5 with decomposer groups-such as microorganisms and insects-contributing to variations in the decomposition rates2,6,7. At the global scale, the contribution of insects to the decomposition of deadwood and carbon release remains poorly understood7. Here we present a field experiment of wood decomposition across 55 forest sites and 6 continents. We find that the deadwood decomposition rates increase with temperature, and the strongest temperature effect is found at high precipitation levels. Precipitation affects the decomposition rates negatively at low temperatures and positively at high temperatures. As a net effect-including the direct consumption by insects and indirect effects through interactions with microorganisms-insects accelerate the decomposition in tropical forests (3.9% median mass loss per year). In temperate and boreal forests, we find weak positive and negative effects with a median mass loss of 0.9 per cent and -0.1 per cent per year, respectively. Furthermore, we apply the experimentally derived decomposition function to a global map of deadwood carbon synthesized from empirical and remote-sensing data, obtaining an estimate of 10.9 ± 3.2 petagram of carbon per year released from deadwood globally, with 93 per cent originating from tropical forests. Globally, the net effect of insects may account for 29 per cent of the carbon flux from deadwood, which suggests a functional importance of insects in the decomposition of deadwood and the carbon cycle.

Linking drought-induced xylem embolism resistance to wood anatomical traits in Neotropical trees.

Drought-induced xylem embolism is considered to be one of the main factors driving mortality in woody plants worldwide. Although several structure-functional mechanisms have been tested to understand the anatomical determinants of embolism resistance, there is a need to study this topic by integrating anatomical data for many species. We combined optical, laser, and transmission electron microscopy to investigate vessel diameter, vessel grouping, and pit membrane ultrastructure for 26 tropical rainforest tree species across three major clades (magnoliids, rosiids, and asteriids). We then related these anatomical observations to previously published data on drought-induced embolism resistance, with phylogenetic analyses. Vessel diameter, vessel grouping, and pit membrane ultrastructure were all predictive of xylem embolism resistance, but with weak predictive power. While pit membrane thickness was a predictive trait when vestured pits were taken into account, the pit membrane diameter-to-thickness ratio suggests a strong importance of the deflection resistance of the pit membrane. However, phylogenetic analyses weakly support adaptive coevolution. Our results emphasize the functional significance of pit membranes for air-seeding in tropical rainforest trees, highlighting also the need to study their mechanical properties due to the link between embolism resistance and pit membrane diameter-to-thickness ratio. Finding support for adaptive coevolution also remains challenging.

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.

Wood specific gravity and anatomy of branches and roots in 113 Amazonian rainforest tree species across environmental gradients.

Wood specific gravity (WSG) is a strong predictor of tree performance across environmental gradients. Yet it remains unclear how anatomical elements linked to different wood functions contribute to variation in WSG in branches and roots across tropical forests. We examined WSG and wood anatomy in white sand, clay terra firme and seasonally flooded forests in French Guiana, spanning broad environmental gradients found throughout Amazonia. We measured 15 traits relating to branches and small woody roots in 113 species representing the 15 most abundant species in each habitat and representative species from seven monophyletic lineages occurring in all habitats. Fiber traits appear to be major determinants of WSG, independent of vessel traits, in branches and roots. Fiber traits and branch and root WSG increased from seasonally flooded species to clay terra firme species and lastly to white sand species. Branch and root wood traits were strongly phylogenetically constrained. Lineages differed in wood design, but exhibited similar variation in wood structure across habitats. We conclude that tropical trees can invest differently in support and transport to respond to environmental conditions. Wind disturbance and drought stress represent significant filters driving tree distribution of Amazonian forests; hence we suggest that biophysical explanations should receive more attention.

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.

From soil to cacao bean: Unravelling the pathways of cadmium translocation in a high Cd accumulating cultivar of Theobroma cacao L.

The research on strategies to reduce cadmium (Cd) accumulation in cacao beans is currently limited by a lack of understanding of the Cd transfer pathways within the cacao tree. Here, we elucidated the transfer of Cd from soil to the nib (seed) in a high Cd accumulating cacao cultivar. Here, we elucidated the transfer of Cd from soil to the nib (seed) in a high Cd accumulating cacao cultivar through Cd stable isotope fractionation, speciation (X-Ray Absorption Spectroscopy), and localization (Laser Ablation Inductively Coupled Plasma Mass Spectrometry). The plant Cd concentrations were 10-28 higher than the topsoil Cd concentrations and increased as placenta< nib< testa< pod husk< root< leaf< branch. The retention of Cd in the roots was low. Light Cd isotopes were retained in the roots whilst heavier Cd isotopes were transported to the shoots (Δ 114/110 Cd shoot-root = 0.27 ± 0.02 ‰ (weighted average ± standard deviation)). Leaf Cd isotopes were heavier than Cd in the branches (Δ 114/110 Cd IF3 leaves-branch = 0.18 ± 0.01 ‰), confirming typical trends observed in annual crops. Nibs and branches were statistically not distinguishable (Δ 114/110 Cd nib-branch = -0.08‰ ± 0.06 ‰), contrary to the leaves and nibs (Δ 114/110 Cd nib-IF3 leaves = -0.25‰ ± 0.05 ‰). These isotope fractionation patterns alluded to a more direct transfer from branches to nibs rather than from leaves to nibs. The largest fraction (57%) of total plant Cd was present in the branches where it was primarily bound to carboxyl-ligands (60-100%) and mainly localized in the phloem rays and phelloderm of the bark. Cadmium in the nibs was mainly bound to oxygen ligands (60-90%), with phytate as the most plausible ligand. The weight of evidence suggested that Cd was transferred like other nutrients from root to shoot and accumulated in the phloem rays and phelloderm of the branches to reduce the transfer to foliage. Finally, the data indicated that the main contribution of nib Cd was from the phloem tissues of the branch rather than from leaf remobilization. This study extended the limited knowledge on Cd accumulation in perennial, woody crops and revealed that the Cd pathways in cacao are markedly different than in annual crops.

Termite sensitivity to temperature affects global wood decay rates.

Deadwood is a large global carbon store with its store size partially determined by biotic decay. Microbial wood decay rates are known to respond to changing temperature and precipitation. Termites are also important decomposers in the tropics but are less well studied. An understanding of their climate sensitivities is needed to estimate climate change effects on wood carbon pools. Using data from 133 sites spanning six continents, we found that termite wood discovery and consumption were highly sensitive to temperature (with decay increasing >6.8 times per 10°C increase in temperature)-even more so than microbes. Termite decay effects were greatest in tropical seasonal forests, tropical savannas, and subtropical deserts. With tropicalization (i.e., warming shifts to tropical climates), termite wood decay will likely increase as termites access more of Earth's surface.

Within-individual variation of trunk and branch xylem density in tropical trees.

PREMISE OF THE STUDY: Wood density correlates with mechanical and physiological strategies of trees and is important for estimating global carbon stocks. Nonetheless, the relationship between branch and trunk xylem density has been poorly explored in neotropical trees. Here, we examine this relationship in trees from French Guiana and its variation among different families and sites, to improve the understanding of wood density in neotropical forests. METHODS: Trunk and branch xylem densities were measured for 1909 trees in seven sites across French Guiana. A major-axis fit was performed to explore their general allometric relationship and its variation among different families and sites. KEY RESULTS: Trunk xylem and branch xylem densities were significantly positively correlated, and their relationship explained 47% of the total variance. Trunk xylem was on average 9% denser than branch xylem. Family-level differences and interactions between family and site accounted for more than 40% of the total variance, whereas differences among sites explained little variation. CONCLUSIONS: Variation in xylem density within individual trees can be substantial, and the relationship between branch xylem and trunk xylem densities varies considerably among families and sites. As such, whole-tree biomass estimates based on nondestructive branch sampling should correct for both taxonomic and environmental factors. Furthermore, detailed estimates of the vertical distribution of wood density within individual trees are needed to determine the extent to which relying solely upon measures of trunk wood density may cause carbon stocks in tropical forests to be overestimated.

Decoupled leaf and stem economics in rain forest trees.

Cross-species analyses of plant functional traits have shed light on factors contributing to differences in performance and distribution, but to date most studies have focused on either leaves or stems. We extend these tissue-specific analyses of functional strategy towards a whole-plant approach by integrating data on functional traits for 13 448 leaves and wood tissues from 4672 trees representing 668 species of Neotropical trees. Strong correlations amongst traits previously defined as the leaf economics spectrum reflect a tradeoff between investments in productive leaves with rapid turnover vs. costly physical leaf structure with a long revenue stream. A second axis of variation, the 'stem economics spectrum', defines a similar tradeoff at the stem level: dense wood vs. high wood water content and thick bark. Most importantly, these two axes are orthogonal, suggesting that tradeoffs operate independently at the leaf and at the stem levels. By simplifying the multivariate ecological strategies of tropical trees into positions along these two spectra, our results provide a basis to improve global vegetation models predicting responses of tropical forests to global change.

Modeling decay rates of dead wood in a neotropical forest.

Variation of dead wood decay rates among tropical trees remains one source of uncertainty in global models of the carbon cycle. Taking advantage of a broad forest plot network surveyed for tree mortality over a 23-year period, we measured the remaining fraction of boles from 367 dead trees from 26 neotropical species widely varying in wood density (0.23-1.24 g cm(-3)) and tree circumference at death time (31.5-272.0 cm). We modeled decay rates within a Bayesian framework assuming a first order differential equation to model the decomposition process and tested for the effects of forest management (selective logging vs. unexploited), of mode of death (standing vs. downed) and of topographical levels (bottomlands vs. hillsides vs. hilltops) on wood decay rates. The general decay model predicts the observed remaining fraction of dead wood (R2 = 60%) with only two biological predictors: tree circumference at death time and wood specific density. Neither selective logging nor local topography had a differential effect on wood decay rates. Including the mode of death into the model revealed that standing dead trees decomposed faster than downed dead trees, but the gain of model accuracy remains rather marginal. Overall, these results suggest that the release of carbon from tropical dead trees to the atmosphere can be simply estimated using tree circumference at death time and wood density.

Search for antifungal compounds from the wood of durable tropical trees.

Research on antifungal compounds from the durable wood from French Guiana Amazonian forest trees highlights the correlation between the activity of their extracts against wood-rotting fungi and human pathogens. The fractionation of an ethyl acetate extract of Sextonia rubra wood led to the isolation of rubrenolide (1) and rubrynolide (2). The potential of compounds 1 and 2 is described through the evaluation of their activity against 16 pathogenic fungi and their cytotoxicity toward NIH-3T3 mammalian fibroblast cells.

A reverse chemical ecology approach to explore wood natural durability.

The natural durability of wood species, defined as their inherent resistance to wood-destroying agents, is a complex phenomenon depending on many biotic and abiotic factors. Besides the presence of recalcitrant polymers, the presence of compounds with antimicrobial properties is known to be important to explain wood durability. Based on the advancement in our understanding of fungal detoxification systems, a reverse chemical ecology approach was proposed to explore wood natural durability using fungal glutathione transferases. A set of six glutathione transferases from the white-rot Trametes versicolor were used as targets to test wood extracts from seventeen French Guiana neotropical species. Fluorescent thermal shift assays quantified interactions between fungal glutathione transferases and these extracts. From these data, a model combining this approach and wood density significantly predicts the wood natural durability of the species tested previously using long-term soil bed tests. Overall, our findings confirm that detoxification systems could be used to explore the chemical environment encountered by wood-decaying fungi and also wood natural durability.

Functional diversity in gravitropic reaction among tropical seedlings in relation to ecological and developmental traits.

Gravitropism is necessary for plants to control the orientation of their axes while they grow in height. In woody plants, stem re-orientations are costly because they are achieved through diameter growth. The functional diversity of gravitropism was studied to check if the mechanisms involved and their efficiency may contribute to the differentiation of height growth strategies between forest tree species at the seedling stage. Seedlings of eight tropical species were grown tilted in a greenhouse, and their up-righting movement and diameter growth were measured over three months. Morphological, anatomical, and biomechanical traits were measured at the end of the survey. Curvature analysis was used to analyse the up-righting response along the stems. Variations in stem curvature depend on diameter growth, size effects, the increase in self-weight, and the efficiency of the gravitropic reaction. A biomechanical model was used to separate these contributions. Results showed that (i) gravitropic movements were based on a common mechanism associated to similar dynamic patterns, (ii) clear differences in efficiency (defined as the change in curvature achieved during an elementary diameter increment for a given stem diameter) existed between species, (iii) the equilibrium angle of the stem and the anatomical characters associated with the efficiency of the reaction also differed between species, and (iv) the differences in gravitropic reaction were related to the light requirements: heliophilic species, compared to more shade-tolerant species, had a larger efficiency and an equilibrium angle closer to vertical. This suggests that traits determining the gravitropic reaction are related to the strategy of light interception and may contribute to the differentiation of ecological strategies promoting the maintenance of biodiversity in tropical rainforests.

Mesoporosity as a new parameter for understanding tension stress generation in trees.

The mechanism for tree orientation in angiosperms is based on the production of high tensile stress on the upper side of the inclined axis. In many species, the stress level is strongly related to the presence of a peculiar layer, called the G-layer, in the fibre cell wall. The structure of the G-layer has recently been described as a hydrogel thanks to N(2) adsorption-desorption isotherms of supercritically dried samples showing a high mesoporosity (pores size from 2-50 nm). This led us to revisit the concept of the G-layer that had been, until now, only described from anatomical observation. Adsorption isotherms of both normal wood and tension wood have been measured on six tropical species. Measurements show that mesoporosity is high in tension wood with a typical thick G-layer while it is much less with a thinner G-layer, sometimes no more than normal wood. The mesoporosity of tension wood species without a G-layer is as low as in normal wood. Not depending on the amount of pores, the pore size distribution is always centred around 6-12 nm. These results suggest that, among species producing fibres with a G-layer, large structural differences of the G-layer exist between species.

Palikur traditional roundwood construction in eastern French Guiana: ethnobotanical and cultural perspectives.

BACKGROUND: Palikur Amerindians live in the eastern part of French Guiana which is undergoing deep-seated changes due to the geographical and economic opening of the region. So far, Palikur's traditional ecological knowledge is poorly documented, apart from medicinal plants. The aim of this study was to document ethnobotanical practices related to traditional construction in the region. METHODS: A combination of qualitative and quantitative methods was used. Thirty-nine Palikur men were interviewed in three localities (Saint-Georges de l'Oyapock, Regina and Trois-Palétuviers) between December 2013 and July 2014. Twenty-four inventories of wood species used in traditional buildings were conducted in the villages, as well as ethnobotanical walks in the neighboring forests, to complete data about usable species and to determine Linnaean names. RESULTS: After an ethnographic description of roundwood Palikur habitat, the in situ wood selection process of Palikur is precisely described. A total of 960 roundwood pieces were inventoried in situ according to Palikur taxonomy, of which 860 were beams and rafters, and 100 posts in 20 permanent and 4 temporary buildings. Twenty-seven folk species were identified. Sixty-three folk species used in construction were recorded during ethnobotanical walks. They correspond to 263 botanical species belonging to 25 families. Posts in permanent buildings were made of yawu (Minquartia guianensis) (51%) and wakap (Vouacapoua americana) (14%). Beams and rafters were made of wood from Annonaceae (79%) and Lecythidaceae (13%) families. The most frequently used species were kuukumwi priye (Oxandra asbeckii), kuukumwi seyne (Pseudoxandra cuspidata), and pukuu (Xylopia nitida and X. cayennensis). CONCLUSIONS: Although the Palikur's relationship with their habitat is undergoing significant changes, knowledge about construction wood is still very much alive in the Oyapock basin. Many people continue to construct traditional buildings alongside modern houses, using a wide array of species described here for the first time, along with the techniques used.

New Insights on Wood Dimensional Stability Influenced by Secondary Metabolites: The Case of a Fast-Growing Tropical Species Bagassa guianensis Aubl.

Challenging evaluation of tropical forest biodiversity requires the reporting of taxonomic diversity but also the systematic characterization of wood properties in order to discover new promising species for timber industry. Among wood properties, the dimensional stability is regarded as a major technological characteristic to validate whether a wood species is adapted to commercial uses. Cell structure and organization are known to influence the drying shrinkage making wood density and microfibrils angle markers of choice to predict wood dimensional stability. On the contrary the role of wood extractive content remains unclear. This work focuses on the fast-growing tropical species Bagassa guianensis and we report herein a correlation between heartwood drying shrinkage and extractive content. Chemical extractions and shrinkage experiments were performed on separate wood twin samples to better evaluate correctly how secondary metabolites influence the wood shrinkage behaviour. Extractive content were qualitatively and quantitatively analysed using HPLC and NMR spectroscopy. We found that B guianensis heartwood has a homogeneous low shrinkage along its radius that could not be explained only by its basic density. In fact the low drying shrinkage is correlated to the high extractive content and a corrected model to improve the prediction of wood dimensional stability is presented. Additionally NMR experiments conducted on sapwood and heartwood extracts demonstrate that secondary metabolites biosynthesis occurs in sapwood thus revealing B. guianensis as a Juglans-Type heartwood formation. This work demonstrates that B. guianensis, a fast-growing species associated with high durability and high dimensional stability, is a good candidate for lumber production and commercial purposes.

The termiticidal activity of Sextonia rubra (Mez) van der Werff (Lauraceae) extract and its active constituent rubrynolide.

BACKGROUND: Termites are degradation agents that inflict severe damage on wood. Some long-lasting Amazonian trees can resist these insects by producing toxic secondary metabolites. These metabolites could potentially replace synthetic termiticidal products which are becoming more restricted to use. RESULTS: Sextonia rubra is resistant to termite-induced degradation. It has been demonstrated that this species naturally produces an ethyl-acetate-soluble termiticidal metabolite, rubrynolide, to protect its wood. Assays in the presence of tropical and invasive termites established that both rubrynolide and crude ethyl acetate extract from S. rubra wood can be used as a treatment for the protection of sensitive woods against termites. CONCLUSION: Rubrynolide and S. rubra extract are promising candidates for the replacement of synthetic termiticides.

Secondary metabolites of Bagassa guianensis Aubl. wood: a study of the chemotaxonomy of the Moraceae family.

In order to explain the durability of the Moraceae plant family, phytochemistry of Bagassa guianensis was performed. Ethyl acetate extract was obtained from the heartwood and 18 secondary metabolites were isolated, including 6 moracins [6-O-methyl-moracin M, 6-O-methyl-moracin N and moracin Z; previously identified: moracin M, moracin N and moracin P], 8 stilbenoids [presently identified: (-)-epialboctalol and arachidin 4; previously identified: alboctalol, trans-resveratrol, arachidin 2, trans-oxyresveratrol and artogomezianol], 3 previously identified flavonoids, steppogenin, katuranin and dihydromorin, beta-sitosterol and resorcinol. Previous studies suggest that stilbenoids are responsible for the natural durability of wood. Our study has determined that B. guianensis is closely related to Morus sp. in phylogeny and should be included in the Moreae sensu stricto tribe of the Moraceae family.

Biomechanics of buttressed trees: bending strains and stresses.

The different hypotheses about buttress function and formation mainly involve mechanical theory. Forces were applied to two trees of Sloanea spp., a tropical genus that develops typical thin buttresses, and the three-dimensional strains were measured at different parts of the trunk base. Risks of failure were greater on the buttress sides, where shear and tangential stresses are greater, not on the ridges, in spite of high longitudinal (parallel to the grain) stresses. A simple beam model, computed from the second moment of area of digitized cross sections, is consistent with longitudinal strain variations but cannot predict accurately variations with height. Patterns of longitudinal strain variation along ridges are very different in the two individuals, owing to a pronounced lateral curvature in one specimen. The constant stress hypothesis is discussed based on these results. Without chronological data during the development of the tree, it cannot be proved that buttress formation is activated by stress or strain.