Nathalie Bréda

Climate-tree-growth relationships of European beech (Fagus sylvatica L.) in the French Permanent Plot Network (RENECOFOR)

The influence of climate on the radial growth of Fagus sylvatica was investigated using 15 chronologies developed from mature stands of the French Permanent Plot Network (RENECOFOR) growing under different climatic and soil conditions. The relationships between climate and ring widths were analyzed using extreme growth years, simple correlations and response functions analysis. Monthly climatic regressors were derived by a physiological water balance model that used daily climatic data and stand parameters to estimate soil water deficits. The three most frequent negative pointer years (1959, 1989, 1976) result from a particularly intense and durable drought, whereas positive years (1977, 1958) coincide with wet conditions. The total ring chronology variance attributable to climate averages 34.1% (15.8% –57%). Current early-summer soil water deficit enters in 10 models and the deficit in June explains alone a large part of the radial growth variability (mean value: 26.6%). Temperature or soil water deficit for the other months and weather conditions during the previous season were of little consistency across stands. The response pattern of earlywood is very similar and the percentage of variance explained is higher (16.2% –57.8%). Latewood widths present a different response pattern. High minimum temperature in August and/or September often favour wide latewood widths and monthly water deficits play a secondary role. The percentage of variance explained ranges from 8.8% to 67.4%. Soil water capacity strongly modulates ring characteristics and climate-growth relationships. Mean sensitivity, expressed population signal, signal-to-noise ratio and the strength of growth-climate correlations increase with decreasing soil water capacity.

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.

Ten years of fluxes and stand growth in a young beech forest at Hesse, North-eastern France

Abstract• Water and carbon fluxes, as measured by eddy covariance, climate, soil water content, leaf area index, tree biomass, biomass increment (BI), litter fall and mortality were monitored for 10 successive years in a young beech stand in Hesse forest (north-eastern France) under contrasting climatic and management conditions.• Large year-to-year variability of net carbon fluxes (NEE) and to a lesser extent, of tree growth was observed. The variability in NEE (coefficient of variation, CV = 44%) was related to both gross primary production (GPP) and to variations in total ecosystem respiration (TER), each term showing similar and lower interannual variability (CV = 14%) than NEE. Variation in the annual GPP was related to: (i) the water deficit duration and intensity cumulated over the growing season, and (ii) the growing season length, i.e. the period of carbon uptake by the stand. Two thinnings occurring during the observation period did not provoke a reduction in either GPP, water fluxes, or in tree growth. Interannual variation of TER could not be explained by any annual climatic variables, or LAI, and only water deficit duration showed a poor correlation. Annual biomass increment was well correlated to water shortage duration and was significantly influenced by drought in the previous year.• The relationship between annual NEE and biomass increment (BI) was poor: in some years, the annual carbon uptake was much higher and in others much lower than tree growth. However this relationship was much stronger and linear (r2 = 0.93) on a weekly to monthly time-scale from budburst to the date of radial growth cessation, indicating a strong link between net carbon uptake and tree growth, while carbon losses by respiration occurring after this date upset this relationship.• Despite the lack of correlation between annual data, the NEE and BI cumulated over the 10 years of observations were very close.• On the annual time-scale, net primary productivity calculated from eddy fluxes and from biological measurements showed a good correlation.Résumé• Les flux d’eau et de dioxyde de carbone, mesurés par la méthode des corrélations turbulentes, le climat, le contenu en eau du sol, l’indice foliaire, la biomasse et l’accroissement en biomasse (BI) des arbres, les chutes de litière et la mortalité ont été suivis en continu pendant 10 années successives dans une jeune hêtraie de la forêt de Hesse (nord-est de la France) en conditions de climat et de gestion contrastées.• Une forte variabilité interannuelle des flux nets de carbone (NEE) et dans une moindre mesure de la croissance des arbres ont été observées. La variabilité de NEE (son coefficient de variation, CV = 44 %) a été mise en relation avec celles de la productivité primaire brute (GPP) et de la respiration totale de l’écosystème (TER), chacun de ces deux termes montrant une variabilité similaire et plus faible (CV = 14 %) que pour NEE. Les variations de la GPP annuelle étaient sous la dépendance : (i) de la durée et de l’intensité du déficit hydrique cumulé sur la saison de végétation, (ii) la longueur de la saison de végétation, définie comme la période où le peuplement absorbe du dioxyde de carbone. Deux éclaircies pratiquées pendant la période de mesures n’ont pas provoqué de réduction ni de GPP, ni du flux d’évapotranspiration, ni de l’accroissement en biomasse du peuplement. Les variations interannuelles de TER n’ont pu être expliquées par aucune des variables climatiques au pas de temps annuel, ni par le LAI, mais seulement par la durée du déficit hydrique du sol, mais avec une corrélation médiocre. L’accroissement annuel en biomasse était fortement corrélé à la durée de la contrainte hydrique de la même année mais aussi influencé de façon significative par la celle de l’année précédente.• La relation entre la NEE annuelle et l’accroissement en biomasse (BI) n’était pas significative : selon les années, l’absorption de carbone par le peuplement était beaucoup plus forte ou plus faible que l’accroissement du peuplement. Toutefois, nous avons observé une corrélation beaucoup plus élevée et linéaire (r2 = 0,93) sur une base de temps hebdomadaire à mensuelle pendant la période allant du débourrement à la date d’arrêt de croissance radiale, ce qui indique un couplage fort entre l’acquisition du carbone et la croissance des arbres, alors que la perte de carbone par respiration en dehors de cette période découple cette relation.• En dépit de l’absence de corrélation entre NEE et BI au pas de temps annuel, le cumul de NEE et celui de BI sur les 10 années ont été très proches.• Par contre, la productivité primaire nette annuelle, calculée à partir des mesures de flux et biométriques sur le peuplement a montré un bon accord.

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.

Modelling variability of wood density in beech as affected by ring age, radial growth and climate

Although it has been recognized as a key parameter of wood quality and a good source of information on growth, annual wood density has been little studied within diffuse-porous trees such as beech ( Fagus sylvatica Liebl.). In this paper we examine the variability encountered in beech ring density series and analyze the influences of ring age, ring width, climate and between-tree variability on density. Thirty ring sequences were sampled from 55-year- old dominant beech trees growing within the same stand; ring density and width were measured using radiography. Ring density proved to be less variable through time than ring width. The relationship between these two variables was less than observed in ring-porous trees and it showed great variation between trees. The sensitivity of ring width and density to climate was also different; width was strongly linked to soil water deficit whereas density was correlated to temperature and August rainfall. Unlike ring width, wood density showed sensitivity towards climatic characteristics of the late growing season. A large part of annual density variability remains unexplained, even using advanced modelled water balance variables. We hypothesize that a significant part of the tree ring is under internal control. We also demonstrated great inter-tree variability (the tree effect) in ring density, which has an influence on density but not on trees’ response to climate.

Seasonal changes of C and N non-structural compounds in the stem sapwood of adult sessile oak and beech trees

We assessed the pools of non-structural nitrogen compounds (NSNC) through a year, thereby addressing the question of whether mature sessile oak [Quercus petraea (Matt.) Liebl.] and beech (Fagus sylvatica L.), which differ in wood anatomy and growth patterns, exhibit contrasting seasonal dynamics of NSNC pools as previously shown for non-structural carbohydrate (NSC) pools. Seasonal fluctuations of NSNC (amino acids and soluble proteins) and NSC (starch and soluble sugars) pools were analyzed in the inner and the outer stem sapwood. In oak, NSC showed marked seasonal variation within the stem sapwood (accumulation during winter and decrease during bud burst and early wood growth), whereas in beech seasonal fluctuations in NSC were of minor amplitude. Even if the distribution and intensity of the NSNC pools differed between the two species, NSNC of the stem sapwood did not show seasonal variation. The most significant change in NSNC pools was the seasonal fluctuation of protein composition. In both species, two polypeptides of 13 kDa (PP13) and 26 kDa (PP26) accumulated during the coldest period in parallel with starch to sugar conversion and disappeared with the onset of spring growth. The absence of seasonal changes in total soluble protein concentration suggests that the polypeptides are involved in the internal nitrogen (N) cycling of the stem rather than in N storage and remobilization to the other growing organs of the tree.

A Retrospective Isotopic Study of Spruce Decline in the Vosges Mountains (France)

The objective of this study was to assess the time variation of mineral and water stress levels across the life of a declining, Mg-deficient, spruce stand, in order to clarify the factors that caused the decline. Since 1985, strong soil acidification linked to a large leaching of nitrate and base cations was measured at the study site. In 1994, 5 trees were felled and tree rings were measured and analysed for Ca, Mg, K, Sr, 13C12C and 87Sr/86Sr isotopic ratios. Strontium pools and fluxes as well as root Sr isotope ratio in relation to depth were also measured. Wood chemical concentrations and isotope ratios were strongly related to the dominance status of each tree. On average during the study period, the 87Sr/86Sr ratio of spruce wood decreased. Using a mechanistic model computing long term variations of 87Sr/86Sr ratio in trees and soils, we reproduced the observed trend by simulating soil acidification – increasing Sr drainage from the whole profile, and particularly from the organic horizon –, and root uptake becoming more superficial with time. Between 1952 and 1976, tree ring Δ 13C decreased strongly and continuously, which, in addition to other factors, might be related to an increase in water stress. Thus, a decrease in rooting depth, possibly related to soil acidification, appeared as a possible cause for the long term increase in water stress. The extreme drought event of 1976 appears to have revealed and triggered the decline.

Douglas-fir is vulnerable to exceptional and recurrent drought episodes and recovers less well on less fertile sites

ContextSince the 2003 drought and heat wave, there have been many reports of Douglas-fir decline and dieback in France. Given the climate change that is predicted, more frequent drought episodes could induce recurrent decline processes. The nature and background of this threat requires a careful assessment.AimsThe objectives of this study were: (1) to test the hypothesis that the major climate hazard that accounts for the decline in Douglas-fir growth is drought, and (2) to identify the main vulnerability factors involved.MethodsDecline and recovery of radial growth were quantified of Douglas-fir in two particularly affected regions with a dendroecological approach; 899 trees were sampled on 58 plots. Mean climate data, ecological and dendrometric stand characteristics were tested in order to identify potential vulnerability factorsResultsA clear relationship was evidenced between growth and soil water deficit. A severe decline of radial growth was induced by the 2003 drought and was maintained during subsequent years due to recurrent drought episodes. Growth recovery at the stand level was enhanced by soil nutrient fertility.ConclusionRadial growth of Douglas-fir is clearly driven by the intensity of drought. In a context of climate change, an adaptive silvicultural strategy is required in order to cope with more frequent drought events. To mitigate the frequency and the intensity of water shortage episodes, site selection must take into account not only local climate characteristic but also soil properties like maximum extractable water content. Soil nitrogen fertility was found to play a key role for an efficient recovery of radial growth after drought episodes and thus must be preserved.

Wood density proxies of adaptive traits linked with resistance to drought in Douglas fir (Pseudotsuga menziesii (Mirb.) Franco)

AbstractKey messageProxies of adaptive traits for resistance to drought were discovered among original annual ring density variables in Douglas fir.Abstract A comparison of dead and surviving Douglas fir trees following the 2003 drought was made to define proxies of adaptive traits for resistance to drought. Increment cores were sampled from trees from three French regions: Centre, Midi-Pyrénées and Burgundy. Original tree-ring variables were calculated, based on a sliding density criterion dividing the microdensity profile into high- and low-density segments. Tree rings were analysed at each site in a number of consecutive annual rings before the 2003 drought event. Comparison between pairs of surviving and dead trees and between pairs of randomly selected trees (whether dead or alive) supports the evidence of systematic dissimilarities between surviving and dead trees in a number of original density variables. Correlation analysis between original and conventional ring density variables indicates a weak association. We found that the surviving trees were denser than the dead trees in all three sites, but that the denser part of the ring varied from region to region. We identified several original density variables intended to be used as proxies of adaptive traits in future studies of genetic determinism of Douglas fir resistance to drought.

Nitrogen sources for current-year shoot growth in 50-year-old sessile oak trees: an in situ 15N labeling approach

We used long-term in situ (15)N labeling of the soil to investigate the contribution of the two main nitrogen (N) sources (N uptake versus N reserves) to sun shoot growth from bud burst to full leaf expansion in 50-year-old sessile oaks. Recovery of (15)N by growing compartments (leaves, twigs and buds) and presence of (15)N in phloem sap were checked weekly. During the first 2 weeks following bud burst, remobilized N contributed ~90% of total N in growing leaves and twigs. Nitrogen uptake from the soil started concomitantly with N remobilization but contributed only slightly to bud burst. However, the fraction of total N due to N uptake increased markedly once bud burst had occurred, reaching 27% in fully expanded leaves and 18% in developed twigs. In phloem sap, the (15)N label appeared a few days after the beginning of labeling and increased until the end of bud burst, and then decreased at full leaf expansion in June. Of all the shoot compartments, leaves attracted most of the absorbed N, which accounted for 68% of new N in shoots, whereas twigs and new buds accounted for only 28 and 3%, respectively. New N allocated to leaves increased from unfolding to full expansion as total N concentration in the leaves decreased. Our results underline the crucial role played by stored N in rapid leaf growth and in the sustained growth of oak trees. Any factors that reduce N storage in autumn may therefore impair spring shoot growth.