Philippe Thaler

Modelling the influence of assimilate availability on root growth and architecture

A model has been designed to simulate rubber seedling root development as related to assimilate availability. Each root of the system is defined both as an element of a network of axes, characterized by its order, position and connections and as an individual sink competing for assimilates. At each time step, the growth of each root is calculated as a function of its own growth potential and of assimilate availability calculated within the whole plant. The potential elongation rate of a root is estimated by its apical diameter, which reflects the size of the meristem. When a root is initiated, the apical diameter depends on root type, but it varies thereafter according to assimilate availability. Thus, the latter controls both current and potential elongation. The model was able to simulate periodicity in root development as related to shoot growth and to reproduce differences in sensitivity to assimilate availability related to root type. It thereby validated the hypothesis that root growth but also root system architecture depend on assimilate allocation and that apical diameter is a good indicator of root growth potential. Provided that specific calibration is done, this model may be used for other species.

Photosynthetic capacity and temperature responses of photosynthesis of rubber trees (Hevea brasiliensis Müll. Arg.) acclimate to changes in ambient temperatures

The aim of this study was to assess the temperature response of photosynthesis in rubber trees (Hevea brasiliensis Müll. Arg.) to provide data for process-based growth modeling, and to test whether photosynthetic capacity and temperature response of photosynthesis acclimates to changes in ambient temperature. Net CO2 assimilation rate (A) was measured in rubber saplings grown in a nursery or in growth chambers at 18 and 28°C. The temperature response of A was measured from 9 to 45°C and the data were fitted to an empirical model. Photosynthetic capacity (maximal carboxylation rate, Vcmax, and maximal light driven electron flux, Jmax) of plants acclimated to 18 and 28°C were estimated by fitting a biochemical photosynthesis model to the CO2 response curves (A–Ci curves) at six temperatures: 15, 22, 28, 32, 36 and 40°C. The optimal temperature for A (Topt) was much lower in plants grown at 18°C compared to 28°C and nursery. Net CO2 assimilation rate at optimal temperature (Aopt), Vcmax and Jmax at a reference temperature of 25°C (Vcmax25 and Jmax25) as well as activation energy of Vcmax and Jmax (EaV and EaJ) decreased in individuals acclimated to 18°C. The optimal temperature for Vcmax and Jmax could not be clearly defined from our response curves, as they always were above 36°C and not far from 40°C. The ratio Jmax25/Vcmax25 was larger in plants acclimated to 18°C. Less nitrogen was present and photosynthetic nitrogen use efficiency (Vcmax25/Na) was smaller in leaves acclimated to 18°C. These results indicate that rubber saplings acclimated their photosynthetic characteristics in response to growth temperature, and that higher temperatures resulted in an enhanced photosynthetic capacity in the leaves, as well as larger activation energy for photosynthesis.

Carbohydrate storage in wood and bark of rubber trees submitted to different level of C demand induced by latex tapping

When the current level of carbohydrates produced by photosynthesis is not enough to meet the C demand for maintenance, growth or metabolism, trees use stored carbohydrates. In rubber trees (Hevea brasiliensis Muell. Arg.), however, a previous study (Silpi U., A. Lacointe, P. Kasemsap, S. Thanisawanyangkura, P. Chantuma, E. Gohet, N. Musigamart, A. Clement, T. Améglio and P. Thaler. 2007. Carbohydrate reserves as a competing sink: evidence from tapping the rubber tree. Tree Physiol. 27:881-889) showed that the additional sink created by latex tapping results not in a decrease, but in an increase in the non-structural carbohydrate (NSC) storage in trunk wood. In this study, the response of NSC storage to latex tapping was further investigated to better understand the trade-off between latex regeneration, biomass and storage. Three tapping systems were compared to the untapped Control for 2 years. Soluble sugars and starch were analyzed in bark and wood on both sides of the trunk, from 50 to 200 cm from the ground. The results confirmed over the 2 years that tapped trees stored more NSC, mainly starch, than untapped Control. Moreover, a double cut alternative tapping system, which produced a higher latex yield than conventional systems, led to even higher NSC concentrations. In all tapped trees, the increase in storage occurred together with a reduction in trunk radial growth. This was interpreted as a shift in carbon allocation toward the creation of reserves, at the expense of growth, to cover the increased risk induced by tapping (repeated wounding and loss of C in latex). Starch was lower in bark than in wood, whereas it was the contrary for soluble sugars. The resulting NSC was twice as low and less variable in bark than in wood. Although latex regeneration occurs in the bark, changes related to latex tapping were more marked in wood than in bark. From seasonal dynamics and differences between the two sides of the trunk in response to tapping, we concluded that starch in wood behaved as the long-term reserve compartment at the whole trunk level, whereas starch in bark was a local buffer. Soluble sugars behaved like an intermediate, ready-to-use compartment in both wood and bark. Finally, the dynamics of carbohydrate reserves appears a relevant parameter to assess the long-term performance of latex tapping systems.

Why are laterals less affected than main axes by homogeneous unfavourable physical conditions? A model-based hypothesis

When plants develop in strong soils, growth of the root system is generally depressed. However, branching and elongation of branches are often less affected than growth of the main axes, whenever the whole root system encounters even-impeded conditions. On the basis of a model simulating root growth and architecture as related to assimilate availability, we propose a simple hypothesis to explain such behaviour. In the model, growth of each root depends on its own elongation potential, which is estimated by its apical diameter. The potential elongation rate–apical diameter relationship is the same for all the roots of the system and is described by a monomolecular function. Our hypothesis is that the effect of soil strength can be simulated by introducing an impedance factor in the definition of root maximum potential elongation rate, common to the whole root system. When such impedance factor is applied, it affects more the potential of larger roots (main axes) than that of thinner roots (secondary and tertiary branches). Simulations provided in high impedance conditions led to root systems characterised by short taproots, whereas growth of secondary roots was unaffected and growth of tertiary roots was enhanced. Actual branching density was also higher, although branching rules have been unchanged. Such simulated systems where similar to that observed in strong soils. Friction laws or pore size can be involved in the larger reduction of the potential growth of main axes. Moreover, when growth of main axes is restricted, assimilate availability becomes higher for branches and that could explain that their growth could be increased in a homogeneous strong soil.

Effect of leaf age and position on light-saturated CO2 assimilation rate, photosynthetic capacity, and stomatal conductance in rubber trees

Shoots of the tropical latex-producing tree Hevea brasiliensis (rubber tree) grow according to a periodic pattern, producing four to five whorls of leaves per year. All leaves in the same whorl were considered to be in the same leaf-age class, in order to assess the evolution of photosynthesis with leaf age in three clones of rubber trees, in a plantation in eastern Thailand. Light-saturated CO2 assimilation rate (Amax) decreased more with leaf age than did photosynthetic capacity (maximal rate of carboxylation, Vcmax, and maximum rate of electron transport, Jmax), which was estimated by fitting a biochemical photosynthesis model to the CO2-response curves. Nitrogen-use efficiency (Amax/Na, Na is nitrogen content per leaf area) decreased also with leaf age, whereas Jmax and Vcmax did not correlate with Na. Although measurements were performed during the rainy season, the leaf gas exchange parameter that showed the best correlation with Amax was stomatal conductance (gs). An asymptotic function was fitted to the Amax-gs relationship, with R2 = 0.85. Amax, Vcmax, Jmax and gs varied more among different whorls in the same clone than among different clones in the same whorl. We concluded that leaf whorl was an appropriate parameter to characterize leaves for the purpose of modelling canopy photosynthesis in field-grown rubber trees, and that stomatal conductance was the most important variable explaining changes in Amax with leaf age in rubber trees.

Differential gene expression in different types of Hevea brasiliensis roots

Three types of roots (taproots, first order laterals and second order laterals) were functionally characterized on 7-month-old in vitro plantlets regenerated by somatic embryogenesis in Hevea brasiliensis. A histological analysis revealed different levels of differentiation depending on root diameter. A primary structure was found in first and second order lateral roots, while taproots displayed a secondary structure. The expression of 48 genes linked to some of the regulatory pathways acting in roots was compared in leaves, stems and the different types of roots by real-time RT-PCR. Thirteen genes were differentially expressed in the different organs studied in plants grown under control conditions. Nine additional other genes were differentially regulated between organs under water deficit conditions. In addition, 10 genes were significantly regulated in response to water deficit, including 8 regulated mainly in lateral roots types. Our results suggest that the regulation of gene expression in lateral roots is different than that in taproots, which have a main role in nutrient uptake and transport, respectively.

Impact of tapping and soil water status on fine root dynamics in a rubber tree plantation in Thailand

Fine roots (FR) play a major role in the water and nutrient uptake of plants and contribute significantly to the carbon and nutrient cycles of ecosystems through their annual production and turnover. FR growth dynamics were studied to understand the endogenous and exogenous factors driving these processes in a 14-year-old plantation of rubber trees located in eastern Thailand. FR dynamics were observed using field rhizotrons from October 2007 to October 2009. This period covered two complete dry seasons (November to March) and two complete rainy seasons (April to October), allowing us to study the effect of rainfall seasonality on FR dynamics. Rainfall and its distribution during the two successive years showed strong differences with 1500 and 950 mm in 2008 and 2009, respectively. FR production (FRP) completely stopped during the dry seasons and resumed quickly after the first rains. During the rainy seasons, FRP and the daily root elongation rate (RER) were highly variable and exhibited strong annual variations with a total FRP of 139.8 and 40.4 mm-2 and an average RER of 0.16 and 0.12 cm day-1 in 2008 and 2009, respectively. The significant positive correlations found between FRP, RER, the appearance of new roots, and rainfall at monthly intervals revealed the impact of rainfall seasonality on FR dynamics. However, the rainfall patterns failed to explain the weekly variations of FR dynamics observed particularly during the rainy seasons. At this time step, FRP, RER, and the appearance of new FR were negatively correlated to the average soil matric potential measured at a depth of between 30 and 60 cm. In addition, our study revealed a significant negative correlation between FR dynamics and the monthly production of dry rubber. Consequently, latex harvesting might disturb carbon dynamics in the whole tree, far beyond the trunk where the tapping was performed. These results exhibit the impact of climatic conditions and tapping system in the carbon budget of rubber plantations.

Increased light-use efficiency sustains net primary productivity of shaded coffee plants in agroforestry system

In agroforestry systems, shade trees strongly affect the physiology of the undergrown crop. However, a major paradigm is that the reduction in absorbed photosynthetically active radiation is, to a certain extent, compensated by an increase in light-use efficiency, thereby reducing the difference in net primary productivity between shaded and non-shaded plants. Due to the large spatial heterogeneity in agroforestry systems and the lack of appropriate tools, the combined effects of such variables have seldom been analysed, even though they may help understand physiological processes underlying yield dynamics. In this study, we monitored net primary productivity, during two years, on scales ranging from individual coffee plants to the entire plot. Absorbed radiation was mapped with a 3D model (MAESPA). Light-use efficiency and net assimilation rate were derived for each coffee plant individually. We found that although irradiance was reduced by 60% below crowns of shade trees, coffee light-use efficiency increased by 50%, leaving net primary productivity fairly stable across all shade levels. Variability of aboveground net primary productivity of coffee plants was caused primarily by the age of the plants and by intraspecific competition among them (drivers usually overlooked in the agroforestry literature) rather than by the presence of shade trees.

A simple framework to analyze water constraints on seasonal transpiration in rubber tree (Hevea brasiliensis) plantations

Climate change and fast extension in climatically suboptimal areas threaten the sustainability of rubber tree cultivation. A simple framework based on reduction factors of potential transpiration was tested to evaluate the water constraints on seasonal transpiration in tropical sub-humid climates, according pedoclimatic conditions. We selected a representative, mature stand in a drought-prone area. Tree transpiration, evaporative demand and soil water availability were measured every day over 15 months. The results showed that basic relationships with evaporative demand, leaf area index and soil water availability were globally supported. However, the implementation of a regulation of transpiration at high evaporative demand whatever soil water availability was necessary to avoid large overestimates of transpiration. The details of regulation were confirmed by the analysis of canopy conductance response to vapor pressure deficit. The final objective of providing hierarchy between the main regulation factors of seasonal and annual transpiration was achieved. In the tested environmental conditions, the impact of atmospheric drought appeared larger importance than soil drought contrary to expectations. Our results support the interest in simple models to provide a first diagnosis of water constraints on transpiration with limited data, and to help decision making toward more sustainable rubber plantations.

Carbon isotope composition of latex does not reflect temporal variations of photosynthetic carbon isotope discrimination in rubber trees (Hevea brasiliensis)

Latex, the cytoplasm of laticiferous cells localized in the inner bark of rubber trees (Hevea brasiliensis Müll. Arg.), is collected by tapping the bark. Following tapping, latex flows out of the trunk and is regenerated, whereas in untapped trees, there is no natural exudation. It is still unknown whether the carbohydrates used for latex regeneration in tapped trees is coming from recent photosynthates or from stored carbohydrates, and in the former case, it is expected that latex carbon isotope composition of tapped trees will vary seasonally, whereas latex isotope composition of untapped trees will be more stable. Temporal variations of carbon isotope composition of trunk latex (δ(13)C-L), leaf soluble compounds (δ(13)C-S) and bulk leaf material (δ(13)C-B) collected from tapped and untapped 20-year-old trees were compared. A marked difference in δ(13)C-L was observed between tapped and untapped trees whatever the season. Trunk latex from tapped trees was more depleted (1.6‰ on average) with more variable δ(13)C values than those of untapped trees. δ(13)C-L was higher and more stable across seasons than δ(13)C-S and δ(13)C-B, with a maximum seasonal difference of 0.7‰ for tapped trees and 0.3‰ for untapped trees. δ(13)C-B was lower in tapped than in untapped trees, increasing from August (middle of the rainy season) to April (end of the dry season). Differences in δ(13)C-L and δ(13)C-B between tapped and untapped trees indicated that tapping affects the metabolism of both laticiferous cells and leaves. The lack of correlation between δ(13)C-L and δ(13)C-S suggests that recent photosynthates are mixed in the large pool of stored carbohydrates that are involved in latex regeneration after tapping.