Meriem Fournier

The use of terrestrial LiDAR technology in forest science: application fields, benefits and challenges

Abstract• IntroductionThe use of terrestrial LiDAR (light detection and ranging) scanners in forest environments is being studied extensively at present due to the high potential of this technology to acquire three-dimensional data on standing trees rapidly and accurately. This article aims to establish the state-of-the-art in this emerging area.• ObjectivesTerrestrial LiDAR has been applied to forest inventory measurements (plot cartography, species recognition, diameter at breast height, tree height, stem density, basal area and plot-level wood volume estimates) and canopy characterisation (virtual projections, gap fraction and three-dimensional foliage distribution). These techniques have been extended to stand value and wood quality assessment. Terrestrial LiDAR also provides new support for ecological applications such as the assessment of the physical properties of leaves, transpiration processes and microhabitat diversity.• ResultsSince 2003, both the capabilities of the devices and data processing technology have improved significantly, with encouraging results. Nevertheless, measurement patterns and device specifications must be selected carefully according to the objectives of the study. Moreover, automated and reliable programmes are still required to process data to make these methodologies applicable specifically to the forest sciences and to fill the gap between time-consuming manual methods and wide-scale remote sensing such as airborne LiDAR scanning.

Woody biomass production lags stem-girth increase by over one month in coniferous forests

Wood is the main terrestrial biotic reservoir for long-term carbon sequestration(1), and its formation in trees consumes around 15% of anthropogenic carbon dioxide emissions each year(2). However, the seasonal dynamics of woody biomass production cannot be quantified from eddy covariance or satellite observations. As such, our understanding of this key carbon cycle component, and its sensitivity to climate, remains limited. Here, we present high-resolution cellular based measurements of wood formation dynamics in three coniferous forest sites in northeastern France, performed over a period of 3 years. We show that stem woody biomass production lags behind stem-girth increase by over 1 month. We also analyse more general phenological observations of xylem tissue formation in Northern Hemisphere forests and find similar time lags in boreal, temperate, subalpine and Mediterranean forests. These time lags question the extension of the equivalence between stem size increase and woody biomass production to intra-annual time scales(3, 4, 5, 6). They also suggest that these two growth processes exhibit differential sensitivities to local environmental conditions. Indeed, in the well-watered French sites the seasonal dynamics of stem-girth increase matched the photoperiod cycle, whereas those of woody biomass production closely followed the seasonal course of temperature. We suggest that forecasted changes in the annual cycle of climatic factors(7) may shift the phase timing of stem size increase and woody biomass production in the future.

Tension wood and opposite wood in 21 tropical rain forest species. 2. Comparison of some anatomical and ultrastructural criteria

The anatomy of tension wood and opposite wood was compared in 21 tropical rain forest trees from 21 species belonging to 18 families from French Guyana. Wood specimens were taken from the upper and lower sides of naturally tilted trees. Measurement of the growth stress level ensured that the two samples were taken from wood tissues in a different mechanical state: highly tensile-stressed wood on the upper side, called tension wood and normally tensile-stressed wood on the lower side, called opposite wood. Quantitative parameters relating to fibres and vessels were measured on transverse sections of both tension and opposite wood to check if certain criteria can easily discriminate the two kinds of wood. We observed a decrease in the frequency of vessels in the tension wood in all the trees studied. Other criteria concerning shape and surface area of the vessels, fibre diameter or cell wall thickness did not reveal any general trend. At the ultrastructural level, we observed that the microfibril angle in the tension wood sample was lower than in opposite wood in all the trees except one (Licania membranacea).

The gravitropic response of poplar trunks: key roles of prestressed wood regulation and the relative kinetics of cambial growth versus wood maturation.

In tree trunks, the motor of gravitropism involves radial growth and differentiation of reaction wood (Archer, 1986). The first aim of this study was to quantify the kinematics of gravitropic response in young poplar (Populus nigra x Populus deltoides, ‘I4551’) by measuring the kinematics of curvature fields along trunks. Three phases were identified, including latency, upward curving, and an anticipative autotropic decurving, which has been overlooked in research on trees. The biological and mechanical bases of these processes were investigated by assessing the biomechanical model of Fournier et al. (1994). Its application at two different time spans of integration made it possible to test hypotheses on maturation, separating the effects of radial growth and cross section size from those of wood prestressing. A significant correlation between trunk curvature and Fourniers model integrated over the growing season was found, but only explained 32% of the total variance. Moreover, over a weeks time period, the model failed due to a clear out phasing of the kinetics of radial growth and curvature that the model does not take into account. This demonstrates a key role of the relative kinetics of radial growth and the maturation process during gravitropism. Moreover, the degree of maturation strains appears to differ in the tension woods produced during the upward curving and decurving phases. Cell wall maturation seems to be regulated to achieve control over the degree of prestressing of tension wood, providing effective control of trunk shape.

Generalized additive models reveal the intrinsic complexity of wood formation dynamics

The intra-annual dynamics of wood formation, which involves the passage of newly produced cells through three successive differentiation phases (division, enlargement, and wall thickening) to reach the final functional mature state, has traditionally been described in conifers as three delayed bell-shaped curves followed by an S-shaped curve. Here the classical view represented by the ‘Gompertz function (GF) approach’ was challenged using two novel approaches based on parametric generalized linear models (GLMs) and ‘data-driven’ generalized additive models (GAMs). These three approaches (GFs, GLMs, and GAMs) were used to describe seasonal changes in cell numbers in each of the xylem differentiation phases and to calculate the timing of cell development in three conifer species [Picea abies (L.), Pinus sylvestris L., and Abies alba Mill.]. GAMs outperformed GFs and GLMs in describing intra-annual wood formation dynamics, showing two left-skewed bell-shaped curves for division and enlargement, and a right-skewed bimodal curve for thickening. Cell residence times progressively decreased through the season for enlargement, whilst increasing late but rapidly for thickening. These patterns match changes in cell anatomical features within a tree ring, which allows the separation of earlywood and latewood into two distinct cell populations. A novel statistical approach is presented which renews our understanding of xylogenesis, a dynamic biological process in which the rate of cell production interplays with cell residence times in each developmental phase to create complex seasonal patterns.

How to determine sapling buckling risk with only a few measurements

Tree buckling risk (actual height/critical buckling height) is an important biomechanical trait of plant growth strategies, and one that contributes to species coexistence. To estimate the diversity of this trait among wide samples, a method that minimizes damage to the plants is necessary. On the basis of the rarely used, complete version of Greenhills model (1881, Proceedings of the Cambridge Philosophical Society 4(2): 65-73), we precisely measured all the necessary parameters on a sample of 236 saplings of 16 species. Then, using sensitivity (variance) analysis, regressions between successive models for risk factors and species ranks and the use of these models on samples of self- and nonself-supporting saplings, we tested different degrees of simplification up to the most simple and widely used formula that assumes that the tree is a cylindrical homogeneous pole. The size factor had the greatest effect on buckling risk, followed by the form factor and the modulus of elasticity of the wood. Therefore, estimates of buckling risk must consider not only the wood properties but especially the form factor. Finally, we proposed a simple but accurate method of assessing tree buckling risk that is applicable to a wide range of samples and that requires mostly nondestructive measurements.

Mapping Radial,Tangential and Longitudinal Shrinkages and Relation to Tension Wood in Discs of the Tropical Tree Symphonia globulifera

Summary A method for measuring shrinkage resulting from drying in the three anisotropic directions is developed and tested. Measurements are performed on sawn discs, a technique which simplifies preparation and enables large numbers of measurements. Shrinkage values can be represented as a map of the disc surface. The results indicate that comparisons between shrinkage distribution and tension wood distribution on the discs show a clear relationship and can be measured with relatively high accuracy in reference to the “shrinkage map”. In the long term, this method could be useful in the timber industry as a means for choosing the direction in which logs are cut depending on their type of wood composition.

Growth and posture control strategies in Fagus sylvatica and Acer pseudoplatanus saplings in response to canopy disturbance

BACKGROUND AND AIMS Forest tree saplings that grow in the understorey undergo frequent changes in their light environment to which they must adapt to ensure their survival and growth. Crown architecture, which plays a critical role in light capture and mechanical stability, is a major component of sapling adaptation to canopy disturbance. Shade-adapted saplings typically have plagiotropic stems and branches. After canopy opening, they need to develop more erect shoots in order to exploit the new light conditions. The objective of this study was to test whether changes in sapling stem inclination occur after canopy opening, and to analyse the morphological changes associated with stem reorientation. METHODS A 4-year canopy-opening field experiment with naturally regenerated Fagus sylvatica and Acer pseudoplatanus saplings was conducted. The appearance of new stem axes, stem basal diameter and inclination along the stem were recorded every year after canopy opening. KEY RESULTS Both species showed considerable stem reorientation resulting primarily from uprighting (more erect) shoot movements in Fagus, and from uprighting movements, shoot elongation and formation of relay shoots in Acer. In both species, the magnitude of shoot uprighting movements was primarily related to initial stem inclination. Both the basal part and the apical part of the stem contributed to uprighting movements. Stem movements did not appear to be limited by stem size or by stem growth. CONCLUSIONS Stem uprighting movements in shade-adapted Fagus and Acer saplings following canopy disturbance were considerable and rapid, suggesting that stem reorientation processes play a significant role in the growth strategy of the species.

Genetic control of stem form in Pinus pinaster Ait. seedlings exposed to lateral light

Abstract In order to compare family differences in stem form determination, seedlings of eight half-sib progenies of Pinus pinaster were exposed to three different light treatments: fixed lateral light (T1), alternate lateral light (T2), and direct sunlight (T3). From spatial coordinates of several points along the stem we computed four variables: global leaning (GL), apical leaning (APL), flexuosity index (FL) and length of the stem (LEN). Based on comparisons of the four variables, the phototropic stimulus caused different responses in stem form at the familial level. Treatment had an important effect on stem form, although the familial response to treatments was quite variable. Heritability values (h2) of the four analyzed variables showed a different pattern of variation for the three treatments, being very stable for LEN and FL, whereas h2 of APL was much greater based on T1 and T2 compared to T3, and h2 of GL based on T2 was much greater compared to T3 and T1. Differences in stem form were due not only to photomorphogenetic response to different light stimuli, but also to the straightening processes such as compression wood formation. The results suggest that lateral light treatments may be useful in revealing differences in efficiency of the mechanisms involved in stem form determination.

Effect of tree size and competition on tension wood production over time in beech plantations and assessing relative gravitropic response with a biomechanical model

PREMISE OF THE STUDY Gravitropic movements are unexpected mechanical processes that could disturb tree design allometries derived from the physics of nonliving bodies. We investigated whether the scaling law of gravitropic performance (power of -2 of stem diameter) derived from integrative biomechanical modeling is disturbed by ontogeny or environment, then discuss the silvicultural and dendroecological consequences. METHODS In a beech (Fagus sylvatica) plantation, four plots with different initial planting densities evolved without any intervention for 26 yr. Regular tree inventories and a silvicultural model were used to monitor competition over time in each plot. The radial production of tension wood was quantified using a cross-section of the stems at 1.30-m height, and an integrative biomechanical model computed the tree gravitropic performance over time. KEY RESULTS All trees developed tension wood over the whole period, with higher amounts at the youngest age, resulting in theoretical lean corrections of ca. 20-30° on the first 4 m of the stem over the whole period. The scaling law of gravitropic performance is slightly larger than the power of -2 of stem diameter. CONCLUSIONS Gravitropic performance in forest ecosystems is mainly limited by size (diameter). Ontogenic acclimation of tension wood formation allows the youngest trees to be more reactive. No additional effect of spacing was found. However, silviculture influences size and, therefore, tree reactivity at a given age. Such results will be helpful for dendroecological approaches that use wood as a marker of environmental disturbances or a trait linked to plant strategies.