A method for the quantification of phototropic and gravitropic sensitivities of plants combining an original experimental device with model-assisted phenotyping: Exploratory test of the method on three hardwood tree species.

Perception of inclination in the gravity field and perception of light direction are two important environmental signals implicated in the control of plant shape and habit. However, their quantitative study in light-grown plants remains a challenge. We present a novel method here to determine the sensitivities to gravitropism and phototropism. The method combines: (i) an original experimental device of isotropic light to disentangle gravitropic and phototropic plant responses; and (ii) model-assisted phenotyping using recent models of tropism perception-the AC model for gravitropism alone and the ArC model for gravitropism combined with phototropism. We first assessed the validity of the AC and ArC models on poplar, the classical species model for woody plants. We then tested the method on three woody species contrasted by their habit and tolerance to shade: poplar (Populus tremula*alba), oak (Quercus petraea) and beech (Fagus sylvatica). The method was found to be effective to quantitatively discriminate the tested species by their ratio of tropistic sensitivities. The method thus appears as an interesting tool to quantitatively determine tropistic sensitivities, a prerequisite for assessing the role of tropisms in the control of the variability of the habit and/or tolerance to shade of woody species in the future.

Temperature heterogeneity over leaf surfaces: the contribution of the lamina microtopography.

Temperature is spatially heterogeneous over leaf surfaces, yet the underlying mechanisms are not fully resolved. We hypothesized that the 3D leaf microtopography determines locally the amount of incoming irradiation flux at leaf surface, thereby driving the temperature gradient over the leaf surface. This hypothesis was tested by developing a model of leaf temperature heterogeneity that includes the development of the leaf boundary layer, the microtopography of the leaf surface and the physiological response of the leaf. Temperature distributions under various irradiation loads (1) over apple leaves based on their 3D microtopography, (2) over simulated flat (2D) apple leaves and (3) over 3D leaves with a transpiration rate distributed as in 2D leaves were simulated. Accuracy of the predictions was quantified by comparing model outputs and thermographic measurements of leaf surface temperature under controlled conditions. Only the model with 3D leaves predicted accurately the spatial heterogeneity of surface temperature over single leaves, whereas the mean temperature was well predicted by both 2D and 3D leaves. We suggest that in these conditions, the 3D leaf microtopography is the primary driver of leaf surface heterogeneity in temperature when the leaf is exposed to a light/heat source.

Mechanical stimuli regulate the allocation of biomass in trees: demonstration with young Prunus avium trees.

BACKGROUND AND AIMS: Plastic tree-shelters are increasingly used to protect tree seedlings against browsing animals and herbicide drifts. The biomass allocation in young seedlings of deciduous trees is highly disturbed by common plastic tree-shelters, resulting in poor root systems and reduced diameter growth of the trunk. The shelters have been improved by creating chimney-effect ventilation with holes drilled at the bottom, resulting in stimulated trunk diameter growth, but the root deficit has remained unchanged. An experiment was set up to elucidate the mechanisms behind the poor root growth of sheltered Prunus avium trees. METHODS: Tree seedlings were grown either in natural windy conditions or in tree-shelters. Mechanical wind stimuli were suppressed in ten unsheltered trees by staking. Mechanical stimuli (bending) of the stem were applied in ten sheltered trees using an original mechanical device. KEY RESULTS: Sheltered trees suffered from poor root growth, but sheltered bent trees largely recovered, showing that mechano-sensing is an important mechanism governing C allocation and the shoot-root balance. The use of a few artificial mechanical stimuli increased the biomass allocation towards the roots, as did natural wind sway. It was demonstrated that there was an acclimation of plants to the imposed strain. CONCLUSIONS: This study suggests that if mechanical stimuli are used to control plant growth, they should be applied at low frequency in order to be most effective. The impact on the functional equilibrium hypothesis that is used in many tree growth models is discussed. The consequence of the lack of mechanical stimuli should be incorporated in tree growth models when applied to environments protected from the wind (e.g. greenhouses, dense forests).

A double-digitising method for building 3D virtual trees with non-planar leaves: application to the morphology and light-capture properties of young beech trees (Fagus sylvatica).

We developed a double-digitising method combining a hand-held electromagnetic digitizer and a non-contact 3D laser scanner. The former was used to record the positions of all leaves in a tree and the orientation angles of their lamina. The latter served to obtain the morphology of the leaves sampled in the tree. As the scanner outputs a cloud of points, software was developed to reconstruct non-planar (NP) leaves composed of triangles, and to compute numerical shape parameters: midrib curvature, torsion and transversal curvature of the lamina. The combination of both methods allowed construction of 3D virtual trees with NP leaves. The method was applied to young beech trees (Fagus sylvatica L.) from different sunlight environments (from 1 to 100% incident light) in a forest in central France. Leaf morphology responded to light availability, with a more bent shape in well-lit leaves. Light interception at the leaf scale by NP leaves decreased from 4 to 10% for shaded and sunlit leaves compared with planar leaves. At the tree scale, light interception by trees made of NP leaves decreased by 1 to 3% for 100% to 1% light, respectively.

Six-year time course of light-use efficiency, carbon gain and growth of beech saplings (Fagus sylvatica) planted under a Scots pine (Pinus sylvestris) shelterwood.

Two-year-old Fagus sylvatica L. saplings were planted under the cover of a Pinus sylvestris L. stand in the French Massif Central. The stand was differentially thinned to obtain a gradient of transmitted photosynthetically active radiation (PAR(t); 0-0.35). Eighteen Fagus saplings were sampled in this gradient, and their growth (basal stem diameter increment) was recorded over six years. Over the same period, morphological parameters (leaf area, number and arrangement in space) were monitored by 3D-digitization. Photosynthetic parameters were estimated with a portable gas-exchange analyzer. Photosynthesis was mainly related to light availability, whereas sapling morphology was mainly driven by sapling size. Annual stem diameter increment was related to the amount of light-intercepting foliage (silhouette to total leaf area ratio (STAR) x total sapling leaf area (LA)) and light availability above the saplings (PAR(t)). However, light-use efficiency, i.e., the slope of the relationship between STAR x LA x PAR(t) and stem diameter increment, decreased over time as a result of a relative decrease in the proportion of photosynthetic tissues to total sapling biomass.

Effect of tapping activity on the dynamics of radial growth of Hevea brasiliensis trees.

Rubber tree (Hevea brasiliensis Müll. Arg.) radial growth dynamics were monitored with displacement sensors, together with latex production, to investigate three aspects of the dual production of latex and wood: (1) the usefulness of fine-scale dendrometric measurements as a physiological tool to detect water shortage through radial growth; (2) the dynamic aspects, both at the seasonal and at the multi-year scale, of the competition between latex and wood production; and (3) the spatial distribution of radial growth rates around the tapping cut. Radial growth of untapped control trees started with the onset of the rainy season and lasted until the onset of the dry season, ceasing completely during the driest period. Displacement sensors provided a sensitive means of detecting water shortage, with a clear correlation between diameter variations and changes in water availability (both daily evapotranspiration and monthly rainfall) over the whole annual cycle. However, the correlation was significantly disturbed in tapped trees. After resumption of tapping, the radial growth rate dropped sharply within two weeks and the effect persisted throughout the whole season, so that the cumulative growth of tapped trees was about half that of untapped trees, with the cumulative growth deficit reaching 80% for the period from mid-June to November. This long-known negative impact of tapping on growth was much stronger in the second year of tapping than in the first, whereas latex production increased significantly between the first and second year of tapping. The increased latex production, which could not be ascribed to climatic conditions, shows that the establishment of an artificial latex sink is a progressive, long-term process likely involving many aspects of metabolism. As expected, ethylene significantly increased latex production in both years; however, ethylene had no effect on the growth rates of tapped trees. Radial growth was differentially affected at different locations around the tapping cut, with growth rates significantly lower in the tapped panel than in the untapped panel, and higher above the cut than below the cut. Thus, caution is needed when deriving whole stem wood production from girth measurements at one location on the stem, especially from girth measurements made close to the tapping cut. This also provides new evidence for the location of the latex regeneration area in the tapped panel, below the cut.

Sudden increase in atmospheric CO2 concentration reveals strong coupling between shoot carbon uptake and root nutrient uptake in young walnut trees.

We studied the short-term (i.e., a few days) effect of a sudden increase in CO2 uptake by shoots on nutrient (NO3-, P ion, K+, Ca2+ and Mg2+) uptake by roots during vegetative growth of young walnut (Juglans nigra x J. major L.) trees. The increase in CO2 uptake was induced by a sudden increase in atmospheric CO2 concentration ([CO2]). Twelve 2-year-old trees were transplanted and grown in perlite-filled pots in a greenhouse. Rates of CO2 uptake and water loss by individual trees were determined by a branch bag method from 3 days before until 6 days after [CO2] was increased. Nutrient uptake rates were measured concurrently by a hydroponic recirculating nutrient solution system that provided non-limiting supplies of water and nutrients. Six control trees were kept in ambient [CO2] (360 ppm), and [CO2] was increased to 550 ppm for one set of three trees and to 800 ppm for another set of three trees. Before imposing the elevated [CO2] treatments, all trees exhibited similar daily water loss, CO2 uptake and nutrient uptake rates when expressed per unit leaf area to account for the tree size effect. Daily water loss rates were only slightly affected by elevated [CO2]. Carbon dioxide uptake rates greatly increased with increasing atmospheric [CO2], and nutrient uptake rates were proportional to CO2 uptake rates during the study period, except for P ion. Our results show that, despite the important carbon and nitrogen storage capacities previously observed in young walnut trees, nutrient uptake by roots is strongly coupled to carbon uptake by shoots over periods of a few days.

Spatial distribution of leaf nitrogen and photosynthetic capacity within the foliage of individual trees: disentangling the effects of local light quality, leaf irradiance, and transpiration.

There is presently no consensus about the factor(s) driving photosynthetic acclimation and the intra-canopy distribution of leaf characteristics under natural conditions. The impact was tested of local (i) light quality (red/far red ratio), (ii) leaf irradiance (PPFD(i)), and (iii) transpiration rate (E) on total non-structural carbohydrates per leaf area (TNC(a)), TNC-free leaf mass-to-area ratio (LMA), total leaf nitrogen per leaf area (N(a)), photosynthetic capacity (maximum carboxylation rate and light-saturated electron transport rate), and leaf N partitioning between carboxylation and bioenergetics within the foliage of young walnut trees grown outdoors. Light environment (quantity and quality) was controlled by placing individual branches under neutral or green screens during spring growth, and air vapour pressure deficit (VPD) was prescribed and leaf transpiration and photosynthesis measured at branch level by a branch bag technique. Under similar levels of leaf irradiance, low air vapour pressure deficit decreased transpiration rate but did not influence leaf characteristics. Close linear relationships were detected between leaf irradiance and leaf N(a), LMA or photosynthetic capacity, and low R/FR ratio decreased leaf N(a), LMA and photosynthetic capacity. Irradiance and R/FR also influenced the partitioning of leaf nitrogen into carboxylation and electron light transport. Thus, local light level and quality are the major factors driving photosynthetic acclimation and intra-canopy distribution of leaf characteristics, whereas local transpiration rate is of less importance.