Eric Dufrêne

Estimation of deciduous forest leaf area index using direct and indirect methods

This study evaluated one semi-direct and three indirect methods for estimating leaf area index (LAI) by comparing these estimates with direct estimates derived from litter collection. The semi-direct method uses a thin metallic needle to count a number of contacts across fresh litter layers. One indirect method is based on the penetration of diffuse global radiation measured over the course of a day. The second indirect method uses the LAI-2000 plant canopy analyser (PCA) which measures diffuse light penetration from five different sky sectors simultaneously. The third indirect method uses the “Demon” portable light sensor to measure the penetration of direct beam sunlight at different zenith angles over the course of half a day. The Poisson model of gap frequency was applied to estimate plant area index (PAI) from observed transmittances using the second and third methods. Litter collection from 11 temperate decidous forests gave values of LAI ranging from 1.7 to 7.5. Estimates based on the needle method showed a significant linear relationship with LAI values obtained from litter collections but were systematically lower (by 6–37%). PAI estimates using all three indirect techniques (fixed light sensor system, LAI-2000 and Demon) showed a strong linear relationship with LAI derived from litter collection. Differences, averaged over all forest stands, between PAI estimates from each of the three indirect methods and LAI from litter collections were below 2%. If we consider that LAI=PAI−WAI (wood area index) then, all three indirect methods underestimated LAI by an additional factor close to the value of WAI. One reason could be a local clumping of architectural canopy components: in particular, the spatial dispositions of branchlets and leaves are not independent, leading to a non-random relationship between the distributions of these two canopy components.

Comparing the intra-annual wood formation of three European species (Fagus sylvatica, Quercus petraea and Pinus sylvestris) as related to leaf phenology and non-structural carbohydrate dynamics

Monitoring cambial phenology and intra-annual growth dynamics is a useful approach for characterizing the tree growth response to climate change. However, there have been few reports concerning intra-annual wood formation in lowland temperate forests with high time resolution, especially for the comparison between deciduous and coniferous species. The main objective of this study was to determine how the timing, duration and rate of radial growth change between species as related to leaf phenology and the dynamics of non-structural carbohydrates (NSC) under the same climatic conditions. We studied two deciduous species, Fagus sylvatica L. and Quercus petraea (Matt.) Liebl., and an evergreen conifer, Pinus sylvestris L. During the 2009 growing season, we weekly monitored (i) the stem radial increment using dendrometers, (ii) the xylem growth using microcoring and (iii) the leaf phenology from direct observations of the tree crowns. The NSC content was also measured in the eight last rings of the stem cores in April, June and August 2009. The leaf phenology, NSC storage and intra-annual growth were clearly different between species, highlighting their contrasting carbon allocation. Beech growth began just after budburst, with a maximal growth rate when the leaves were mature and variations in the NSC content were low. Thus, beech radial growth seemed highly dependent on leaf photosynthesis. For oak, earlywood quickly developed before budburst, which probably led to the starch decrease quantified in the stem from April to June. For pine, growth began before the needles unfolding and the lack of NSC decrease during the growing season suggested that the substrates for radial growth were new assimilates of the needles from the previous year. Only for oak, the pattern determined from the intra-annual growth measured using microcoring differed from the pattern determined from dendrometer data. For all species, the ring width was significantly influenced by growth duration and not by growth rate, which differs from previous studies. The observed between-species difference at the intra-annual scale is key information for anticipating suitability of future species in temperate forests.

Climate change impacts on tree ranges: model intercomparison facilitates understanding and quantification of uncertainty

Model-based projections of shifts in tree species range due to climate change are becoming an important decision support tool for forest management. However, poorly evaluated sources of uncertainty require more scrutiny before relying heavily on models for decision-making. We evaluated uncertainty arising from differences in model formulations of tree response to climate change based on a rigorous intercomparison of projections of tree distributions in France. We compared eight models ranging from niche-based to process-based models. On average, models project large range contractions of temperate tree species in lowlands due to climate change. There was substantial disagreement between models for temperate broadleaf deciduous tree species, but differences in the capacity of models to account for rising CO(2) impacts explained much of the disagreement. There was good quantitative agreement among models concerning the range contractions for Scots pine. For the dominant Mediterranean tree species, Holm oak, all models foresee substantial range expansion.

Age-related variation in carbon allocation at tree and stand scales in beech (Fagus sylvatica L.) and sessile oak (Quercus petraea (Matt.) Liebl.) using a chronosequence approach

Two types of physiological mechanisms can contribute to growth decline with age: (i) the mechanisms leading to the reduction of carbon assimilation (input) and (ii) those leading to modification of the resource economy. Surprisingly, the processes relating to carbon allocation have been little investigated as compared to research on the processes governing carbon assimilation. The objective of this paper was thus to test the hypothesis that growth decrease related to age is accompanied by changes in carbon allocation to the benefit of storage and reproductive functions in two contrasting broad-leaved species: beech (Fagus sylvatica L.) and sessile oak (Quercus petraea (Matt.) Liebl.). Age-related changes in carbon allocation were studied using a chronosequence approach. Chronosequences, each consisting of several even-aged stands ranging from 14 to 175 years old for beech and from 30 to 134 years old for sessile oak, were divided into five or six age classes. In this study, carbon allocations to growth, storage and reproduction were defined as the relative amount of carbon invested in biomass increment, carbohydrate increment and seed production, respectively. Tree-ring width and allometric relationships were used to assess biomass increment at the tree and stand scales. Below-ground biomass was assessed using a specific allometric relationship between root:shoot ratio and age, established from the literature review. Seasonal variations of carbohydrate concentrations were used to assess carbon allocation to storage. Reproduction effort was quantified for beech stands by collecting seed and cupule production. Age-related flagging of biomass productivity was assessed at the tree and stand scales, and carbohydrate quantities in trees increased with age for both species. Seed and cupule production increased with stand age in beech from 56 gC m(-)(2) year(-1) at 30 years old to 129 gC m(-2) year(-1) at 138 years old. In beech, carbon allocation to storage and reproductive functions increased with age to the detriment of carbon allocation to growth functions. In contrast, the carbon balance between growth and storage remained constant between age classes in sessile oak. The contrasting age-related changes in carbon allocation between beech and sessile oak are discussed with reference to the differences in growing environment, phenology and hydraulic properties of ring-porous and diffuse-porous species.

Estimation of forest leaf area index from SPOT imagery using NDVI distribution over forest stands

Leaf area index (LAI) is a key parameter of atmosphere–vegetation exchanges, affecting the net ecosystem exchange and the productivity. At regional or continental scales, LAI can be estimated by remotely‐sensed spectral vegetation indices (SVI). Nevertheless, relationships between LAI and SVI show saturation for LAI values greater than 3–5. This is one of the principal limitations of remote sensing of LAI in forest canopies. In this article, a new approach is developed to determine LAI from the spatial variability of radiometric data. To test this method, in situ measurements for LAI of 40 stands, with three dominant species (European beech, oak and Scots pine) were available over 5 years in the Fontainebleau forest near Paris. If all years and all species are pooled, a good linear relationship without saturation is founded between average stand LAI measurements and a model combining the logarithm of the standard deviation and the skewness of the normalized difference vegetation index (NDVI) (R 2 = 0.73 rmse = 1.08). We demonstrate that this relation can be slightly improved by using different linear models for each year and each species (R 2 = 0.82 rmse = 0.86), but the standard deviation is less sensitive to the species and the year effects than the mean NDVI and is therefore a performing index.

Soil (N) modulates soil C cycling in CO2-fumigated tree stands: a meta-analysis

Under elevated atmospheric CO(2) concentrations, soil carbon (C) inputs are typically enhanced, suggesting larger soil C sequestration potential. However, soil C losses also increase and progressive nitrogen (N) limitation to plant growth may reduce the CO(2) effect on soil C inputs with time. We compiled a data set from 131 manipulation experiments, and used meta-analysis to test the hypotheses that: (1) elevated atmospheric CO(2) stimulates soil C inputs more than C losses, resulting in increasing soil C stocks; and (2) that these responses are modulated by N. Our results confirm that elevated CO(2) induces a C allocation shift towards below-ground biomass compartments. However, the increased soil C inputs were offset by increased heterotrophic respiration (Rh), such that soil C content was not affected by elevated CO(2). Soil N concentration strongly interacted with CO(2) fumigation: the effect of elevated CO(2) on fine root biomass and -production and on microbial activity increased with increasing soil N concentration, while the effect on soil C content decreased with increasing soil N concentration. These results suggest that both plant growth and microbial activity responses to elevated CO(2) are modulated by N availability, and that it is essential to account for soil N concentration in C cycling analyses.

Relationship between photochemical reflectance index and leaf ecophysiological and biochemical parameters under two different water statuses: towards a rapid and efficient correction method using real-time measurements

The use of the photochemical reflectance index (PRI) as a promising proxy of light use efficiency (LUE) has been extensively studied, and some issues have been identified, notably the sensitivity of PRI to leaf pigment composition and the variability in PRI response to LUE because of stress. In this study, we introduce a method that enables us to track the short-term PRI response to LUE changes because of photosynthetically active radiation (PAR) changes. The analysis of these short-term relationships between PRI and LUE throughout the growing season in two species (Quercus robur L. and Fagus sylvatica L.) under two different soil water statuses showed a clear change in PRI response to LUE, which is related to leaf pigment content. The use of an estimated or approximated PRI0, defined as the PRI of perfectly dark-adapted leaves, allowed us to separate the PRI variability due to leaf pigment content changes and the physiologically related PRI variability over both daily (PAR-related) and seasonal (soil water content-related) scales. The corrected PRI obtained by subtracting PRI0 from the PRI measurements showed a good correlation with the LUE over both of the species, soil water statuses and over the entire growing season.

Comparison of seasonal variations in water-use efficiency calculated from the carbon isotope composition of tree rings and flux data in a temperate forest

Tree-ring δ(13) C is often interpreted in terms of intrinsic water-use efficiency (WUE) using a carbon isotope discrimination model established at the leaf level. We examined whether intra-ring δ(13) C could be used to assess variations in intrinsic WUE (W(g), the ratio of carbon assimilation and stomatal conductance to water) and variations in ecosystem WUE (W(t) , the ratio of C assimilation and transpiration) at a seasonal scale. Intra-ring δ(13) C was measured in 30- to 60-µm-thick slices in eight oak trees (Quercus petraea). Canopy W(g) was simulated using a physiologically process-based model. High between-tree variability was observed in the seasonal variations of intra-ring δ(13) C. Six trees showed significant positive correlations between W(g) calculated from intra-ring δ(13) C and canopy W(g) averaged over several days during latewood formation. These results suggest that latewood is a seasonal recorder of W(g) trends, with a temporal lag corresponding to the mixing time of sugars in the phloem. These six trees also showed significant negative correlations between photosynthetic discrimination Δ calculated from intra-ring δ(13) C, and ecosystem W(t), during latewood formation. Despite the observed between-tree variability, these results indicate that intra-ring δ(13) C can be used to access seasonal variations in past W(t).

Deconvolution of pigment and physiologically related photochemical reflectance index variability at the canopy scale over an entire growing season

The sensitivity of the photochemical reflectance index (PRI) to leaf pigmentation and its impacts on its potential as a proxy for light-use efficiency (LUE) have recently been shown to be problematic at the leaf scale. Most leaf-to-leaf and seasonal variability can be explained by such a confounding effect. This study relies on the analysis of PRI light curves that were generated at the canopy scale under natural conditions to derive a precise deconvolution of pigment-related and physiologically related variability in the PRI. These sources of variability were explained by measured or estimated physiologically relevant variables, such as soil water content, that can be used as indicators of water availability and canopy chlorophyll content. The PRI mainly reflected the variability in the pigment content of the canopy. However, the corrected PRI, which was obtained by subtracting the pigment-related seasonal variability from the PRI measurement, was highly correlated with the upscaled LUE measurements. Moreover, the sensitivity of the PRI to the leaf pigment content may mask the PRI versus LUE relationship or result in an artificial relationship that reflects the relationship of chlorophyll versus LUE, depending on the species phenology.

Root exclusion through trenching does not affect the isotopic composition of soil CO2 efflux

Disentangling the autotrophic and heterotrophic components of soil CO2 efflux is critical to understanding the role of soil system in terrestrial carbon (C) cycling. In this study, we combined a stable C-isotope natural abundance approach with the trenched plot method to determine if root exclusion significantly affected the isotopic composition (δ13C) of soil CO2 efflux (RS). This study was performed in different forest ecosystems: a tropical rainforest and two temperate broadleaved forests, where trenched plots had previously been installed. At each site, RS and its δ13C (δ13CRs) tended to be lower in trenched plots than in control plots. Contrary to RS, δ13CRs differences were not significant. This observation is consistent with the small differences in δ13C measured on organic matter from root, litter and soil. The lack of an effect on δ13CRs by root exclusion could be from the small difference in δ13C between autotrophic and heterotrophic soil respirations, but further investigations are needed because of potential artefacts associated with the root exclusion technique.