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Audigeos, D., Brousseau, L., Traissac, S., Scotti-Saintagne, C., & Scotti, I. (2013). Molecular divergence in tropical tree populations occupying environmental mosaics. Journal of Evolutionary Biology, 26(3), 529–544.
Abstract: Unveiling the genetic basis of local adaptation to environmental variation is a major goal in molecular ecology. In rugged landscapes characterized by environmental mosaics, living populations and communities can experience steep ecological gradients over very short geographical distances. In lowland tropical forests, interspecific divergence in edaphic specialization (for seasonally flooded bottomlands and seasonally dry terra firme soils) has been proven by ecological studies on adaptive traits. Some species are nevertheless capable of covering the entire span of the gradient; intraspecific variation for adaptation to contrasting conditions may explain the distribution of such ecological generalists. We investigated whether local divergence happens at small spatial scales in two stands of Eperua falcata (Fabaceae), a widespread tree species of the Guiana Shield. We investigated Single Nucleotide Polymorphisms (SNP) and sequence divergence as well as spatial genetic structure (SGS) at four genes putatively involved in stress response and three genes with unknown function. Significant genetic differentiation was observed among sub-populations within stands, and eight SNP loci showed patterns compatible with disruptive selection. SGS analysis showed genetic turnover along the gradients at three loci, and at least one haplotype was found to be in repulsion with one habitat. Taken together, these results suggest genetic differentiation at small spatial scale in spite of gene flow. We hypothesize that heterogeneous environments may cause molecular divergence, possibly associated to local adaptation in E. falcata. © 2012 European Society For Evolutionary Biology.
Keywords: Candidate genes; Drought; Eperua falcata; Flooding; Neotropics; Outlier loci; Tree genetics
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Malé, P. - J. G., Leroy, C., Humblot, P., Dejean, A., Quilichini, A., & Orivel, J. (2016). Limited gene dispersal and spatial genetic structure as stabilizing factors in an ant-plant mutualism. J. Evol. Biol., 29(12), 2519–2529.
Abstract: Comparative studies of the population genetics of closely associated species are necessary to properly understand the evolution of these relationships because gene flow between populations affects the partners' evolutionary potential at the local scale. As a consequence (at least for antagonistic interactions), asymmetries in the strength of the genetic structures of the partner populations can result in one partner having a co-evolutionary advantage. Here, we assess the population genetic structure of partners engaged in a species-specific and obligatory mutualism: the Neotropical ant-plant, Hirtella physophora, and its ant associate, Allomerus decemarticulatus. Although the ant cannot complete its life cycle elsewhere than on H. physophora and the plant cannot live for long without the protection provided by A. decemarticulatus, these species also have antagonistic interactions: the ants have been shown to benefit from castrating their host plant and the plant is able to retaliate against too virulent ant colonies. We found similar short dispersal distances for both partners, resulting in the local transmission of the association and, thus, inbred populations in which too virulent castrating ants face the risk of local extinction due to the absence of H. physophora offspring. On the other hand, we show that the plant populations probably experienced greater gene flow than did the ant populations, thus enhancing the evolutionary potential of the plants. We conclude that such levels of spatial structure in the partners' populations can increase the stability of the mutualistic relationship. Indeed, the local transmission of the association enables partial alignments of the partners' interests, and population connectivity allows the plant retaliation mechanisms to be locally adapted to the castration behaviour of their symbionts.
Keywords: gene flow; local adaptation; metapopulation; myrmecophyte; population genetics
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Seymour, R. S., White, C. R., & Gibernau, M. (2009). Endothermy of dynastine scarab beetles (Cyclocephala colasi) associated with pollination biology of a thermogenic arum lily (Philodendron solimoesense). J. Exp. Biol., 212(18), 2960–2968.
Abstract: Cyclocephala colasi beetles are facultative endotherms that spend most of their adult lives inside the inflorescences of Philodendron solimoesense, where ambient temperature (T-a) averages about 28 degrees C due to floral thermogenesis. Measurements of respiration within a range of T-a showed that active beetles became spontaneously endothermic at T-a below 28 degrees C but were rarely endothermic above it. There was no evidence of endothermy within the inflorescences, indicating that activities in the floral chamber can occur without the high energy expense of endothermy. Bouts of endothermy occurred at lower T-a in respirometer chambers mainly in the evening, when the insects normally fly from one inflorescence to another, and during the night, when they normally eat and mate within the inflorescence. Patterns of endothermy in individual episodes were studied in non-flying beetles with respirometry and infrared thermal imaging. Heat was generated in the thorax by oscillatory waves of respiration that were coupled with thoracic temperature (T-th) increases. Stationary beetles could regulate T-th at about 33 degrees C independently of T-a between 16 and 29 degrees C. At T-a=20 degrees C, this represents a 116-fold increase in metabolic rate over resting, ectothermic values. Endothermy was clearly a requirement for flight, and beetles departing inflorescences warmed to about 30 degrees C before take-off. During flight, T-th was dependent on T-a, decreasing from 37 to 28 degrees C at T-a of 37 to 20 degrees C, respectively. The lowest T-a at which flight could occur was about 20 degrees C. Thermal conductance of stationary, endothermic beetles increased at higher metabolic rates, probably because of increased ventilatory heat loss.
Keywords: beetle; endothermy; pollination biology; Cyclocephala; Philodendron
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Almeras, T., Derycke, M., Jaouen, G., Beauchene, J., & Fournier, M. (2009). Functional diversity in gravitropic reaction among tropical seedlings in relation to ecological and developmental traits. J. Exp. Bot., 60(15), 4397–4410.
Abstract: 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.
Keywords: Biomechanics; French Guiana; functional diversity; gravitropism; reaction wood; tropical rainforest
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Chang, S. S., Clair, B., Ruelle, J., Beauchene, J., Di Renzo, F., Quignard, F., et al. (2009). Mesoporosity as a new parameter for understanding tension stress generation in trees. J. Exp. Bot., 60(11), 3023–3030.
Abstract: 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.
Keywords: Growth stress; hydrogel; mesoporosity; tension wood
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Coste, S., Roggy, J. C., Schimann, H., Epron, D., & Dreyer, E. (2011). A cost-benefit analysis of acclimation to low irradiance in tropical rainforest tree seedlings: leaf life span and payback time for leaf deployment. J. Exp. Bot., 62(11), 3941–3955.
Abstract: The maintenance in the long run of a positive carbon balance under very low irradiance is a prerequisite for survival of tree seedlings below the canopy or in small gaps in a tropical rainforest. To provide a quantitative basis for this assumption, experiments were carried out to determine whether construction cost (CC) and payback time for leaves and support structures, as well as leaf life span (i) differ among species and (ii) display an irradiance-elicited plasticity. Experiments were also conducted to determine whether leaf life span correlates to CC and payback time and is close to the optimal longevity derived from an optimization model. Saplings from 13 tropical tree species were grown under three levels of irradiance. Specific-CC was computed, as well as CC scaled to leaf area at the metamer level. Photosynthesis was recorded over the leaf life span. Payback time was derived from CC and a simple photosynthesis model. Specific-CC displayed only little interspecific variability and irradiance-elicited plasticity, in contrast to CC scaled to leaf area. Leaf life span ranged from 4 months to > 26 months among species, and was longest in seedlings grown under lowest irradiance. It was always much longer than payback time, even under the lowest irradiance. Leaves were shed when their photosynthesis had reached very low values, in contrast to what was predicted by an optimality model. The species ranking for the different traits was stable across irradiance treatments. The two pioneer species always displayed the smallest CC, leaf life span, and payback time. All species displayed a similar large irradiance-elicited plasticity.
Keywords: Carbon balance; construction cost; functional diversity; leaf life span; payback time; photosynthesis; tropical rainforest
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Christensen-Dalsgaard, K. K., Ennos, A. R., & Fournier, M. (2007). Changes in hydraulic conductivity, mechanical properties, and density reflecting the fall in strain along the lateral roots of two species of tropical trees. J. Exp. Bot., 58(15-16), 4095–4105.
Abstract: Roots have been described as having larger vessels and so greater hydraulic efficiency than the stem. Differences in the strength and stiffness of the tissue within the root system itself are thought to be an adaptation to the loading conditions experienced by the roots and to be related to differences in density. It is not known how potential mechanical adaptations may affect the hydraulic properties of the roots. The change in strength, stiffness, conductivity, density, sapwood area, and second moment of area distally along the lateral roots of two tropical tree species in which the strain is known to decrease rapidly was studied and the values were compared with those of the trunk. It was found that as the strain fell distally along the roots, so did the strength and stiffness of the tissue, whereas the conductivity increased exponentially. These changes appeared to be related to differences in density. In contrast to the distal-most roots, the tissue of the proximal roots had a lower conductivity and higher strength than that of the trunk. This suggests that mechanical requirements on the structure rather than the water potential gradient from roots to branches are responsible for the general pattern that roots have larger vessels than the stem. In spite of their increased transectional area, the buttressed proximal roots were subjected to higher levels of stress and had a lower total conductivity than the rest of the root system. © 2007 The Author(s).
Keywords: Buttress roots; Density; Hydraulic conductivity; Hydraulic-mechanical trade-offs; Modulus of elasticity; Tropical trees; Wood; Elastic moduli; Hydraulic conductivity; Wood; Buttress roots; Hydraulic-mechanical trade-offs; Tropical trees; Forestry; water; article; biomechanics; histology; legume; physiology; plant root; plant stem; tree; wood; Xylopia; Biomechanics; Fabaceae; Plant Roots; Plant Stems; Trees; Water; Wood; Xylopia; Conductivity; Elastic Strength; Forestry; Wood
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Fournier, M., Dlouhá, J., Jaouen, G., & Almeras, T. (2013). Integrative biomechanics for tree ecology: Beyond wood density and strength. J. Exp. Bot., 64(15), 4793–4815.
Abstract: Functional ecology has long considered the support function as important, but its biomechanical complexity is only just being elucidated. We show here that it can be described on the basis of four biomechanical traits, two safety traits against winds and self-buckling, and two motricity traits involved in sustaining an upright position, tropic motion velocity (MV) and posture control (PC). All these traits are integrated at the tree scale, combining tree size and shape together with wood properties. The assumption of trait constancy has been used to derive allometric scaling laws, but it was more recently found that observing their variations among environments and functional groups, or during ontogeny, provides more insights into adaptive syndromes of tree shape and wood properties. However, oversimpli-fed expressions have often been used, possibly concealing key adaptive drivers. An extreme case of oversimplification is the use of wood basic density as a proxy for safety. Actually, as wood density is involved in stiffiness, loads, and construction costs, the impact of its variations on safety is non-trivial. Moreover, other wood features, especially the microfibril angle (MFA), are also involved. Furthermore, wood is not only stiff and strong, but it also acts as a motor for MV and PC. The relevant wood trait for this is maturation strain asymmetry. Maturation strains vary with cell-wall characteristics such as MFA, rather than with wood density. Finally, the need for further studies about the ecological relevance of branching patterns, motricity traits, and growth responses to mechanical loads is discussed. © The Author 2013.
Keywords: Biomechanics; Ecological strategy; Gravitropism; Shape; Size; Trees; Wood
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Levionnois, S., Salmon, C., Alméras, T., Clair, B., Ziegler, C., Coste, S., et al. (2021). Anatomies, vascular architectures, and mechanics underlying the leaf size-stem size spectrum in 42 Neotropical tree species. Journal of Experimental Botany, 72(22), 7957–7969.
Abstract: The leaf size-stem size spectrum is one of the main dimensions of plant ecological strategies. Yet the anatomical, mechanical, and hydraulic implications of small vs. large shoots are still poorly understood. We investigated 42 tropical rainforest tree species in French Guiana, with a wide range of leaf areas at the shoot level. We quantified the scaling of hydraulic and mechanical constraints with shoot size estimated as the water potential difference ΔΨ and the bending angle ΔΦ, respectively. We investigated how anatomical tissue area, flexural stiffness and xylem vascular architecture affect such scaling by deviating (or not) from theoretical isometry with shoot size variation. Vessel diameter and conductive path length were found to be allometrically related to shoot size, thereby explaining the independence between ΔΨ and shoot size. Leaf mass per area, stem length, and the modulus of elasticity were allometrically related with shoot size, explaining the independence between ΔΦ and shoot size. Our study also shows that the maintenance of both water supply and mechanical stability across the shoot size range are not in conflict.
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Verbeeck, H., Peylin, P., Bacour, C., Bonal, D., Steppe, K., & Ciais, P. (2011). Seasonal patterns of CO2 fluxes in Amazon forests: Fusion of eddy covariance data and the ORCHIDEE model. J. Geophys. Res.-Biogeosci., 116(2), G02018.
Abstract: [1] In some regions of the Amazon, global biogeophysical models have difficulties in reproducing measured seasonal patterns of net ecosystem exchange (NEE) of carbon dioxide. The global process-based biosphere model Organizing Carbon and Hydrology in Dynamic Ecosystems (ORCHIDEE) used in this study showed that a standard model parameterization produces seasonal NEE patterns that are opposite in phase to the eddy flux data of the tropical evergreen forest at the Tapajos km 67 site (Brazil), like many other global models. However, we optimized several key parameters of ORCHIDEE using eddy covariance data of the Tapajos km 67 site in order to identify the driving factors of the seasonal variations in CO2 flux in this tropical forest ecosystem. The validity of the retrieved parameter values was evaluated for two other flux tower sites in the Amazon. The different tested optimization scenarios showed that only a few parameters substantially improve the fit to NEE and latent heat data. Our results confirm that these forests have the ability to maintain high transpiration and photosynthesis during the dry season in association with a large soil depth (D-soil = 10 m) and a rooting system density that decreases almost linearly with depth (H-root = 0.1). Previous analyses of seasonal variations in eddy covariance fluxes indicated that higher GPP levels were reached in the dry season compared to the wet season. Our optimization analysis suggests that this pattern could be caused by a leaf flush at the start of the dry season increasing the photosynthetic capacity of the canopy. Nevertheless, the current model structure is not yet able to simulate such a leaf flush, and we therefore suggest improving the ORCHIDEE model by including a specific phenology module that is driven by light availability for the tropical evergreen plant functional types. In addition, our results highlight both the potential and the limitations of flux data to improve global terrestrial models. Several parameters were not identifiable, and the risk of overfitting of the model was illustrated. Nevertheless, we conclude that these models can be improved substantially by assimilating site level flux data over the tropics.
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