Jean-Marc Guehl

Interseasonal comparison of CO2 concentrations, isotopic composition, and carbon dynamics in an Amazonian rainforest (French Guiana)

Abstract Canopy CO2 concentrations in a tropical rainforest in French Guiana were measured continuously for 5 days during the 1994 dry season and the 1995 wet season. Carbon dioxide concentrations ([CO2]) throughout the canopy (0.02–38 m) showed a distinct daily pattern, were well-stratified and decreased with increasing height into the canopy. During both seasons, daytime [CO2] in the upper and middle canopy decreased on average 7–10 μmol mol−1 below tropospheric baseline values measured at Barbados. Within the main part of the canopy (≥ 0.7 m), [CO2] did not differ between the wet and dry seasons. In contrast, [CO2] below 0.7 m were generally higher during the dry season, resulting in larger [CO2] gradients. Supporting this observation, soil CO2 efflux was on average higher during the dry season than during the wet season, either due to diffusive limitations and/or to oxygen deficiency of root and microbial respiration. Soil respiration rates decreased by 40% after strong rain events, resulting in a rapid decrease in canopy [CO2] immediately above the forest floor of about 50␣μmol mol−1. Temporal and spatial variations in [CO2]canopy were reflected in changes of δ13Ccanopy and δ18Ocanopy values. Tight relationships were observed between δ13C and δ18O of canopy CO2 during both seasons (r2 > 0.86). The most depleted δ13Ccanopy and δ18Ocanopy values were measured immediately above the forest floor (δ13C = −16.4‰; δ18O = 39.1‰ SMOW). Gradients in the isotope ratios of CO2 between the top of the canopy and the forest floor ranged between 2.0‰ and 6.3‰ for δ13C, and between 1.0‰ and 3.5‰ for δ18O. The δ13Cleaf and calculated ci/ca of foliage at three different positions were similar for the dry and wet seasons indicating that the canopy maintained a constant ratio of photosynthesis to stomatal conductance. About 20% of the differences in δ13Cleaf within the canopy was accounted for by source air effects, the remaining 80% must be due to changes in ci/ca. Plotting 1/[CO2] vs. the corresponding δ13C ratios resulted in very tight, linear relationships (r2 = 0.99), with no significant differences between the two seasons, suggesting negligible seasonal variability in turbulent mixing relative to ecosystem gas exchange. The intercepts of these relationships that should be indicative of the δ13C of respired sources were close to the measured δ13C of soil respired CO2 and to the δ13C of litter and soil organic matter. Estimates of carbon isotope discrimination of the entire ecosystem, Δe, were calculated as 20.3‰ during the dry season and as 20.5‰ during the wet season.

Responses of antioxidative systems to drought stress in pendunculate oak and maritime pine as modulated by elevated CO2

The aim of the present study was to investigate the effects of an enhanced CO2 concentration alone or in combination with drought stress on antioxidative systems of a deciduous (oak; Quercus robur) and an evergreen (pine; Pinus pinaster) tree species. The seedlings were grown for one season in a greenhouse in tunnels supplied with 350 or 700 [mu]L L-1 CO2. The experiment was repeated in a second year. Antioxidants, protective enzymes, soluble protein, and pigments showed considerable fluctuations in different years. Elevated CO2 caused significant reductions in the activities of superoxide dismutases in both oak and pine. The activities of ascorbate peroxidase and catalase were also reduced in most cases. The activities of dehydroascorbate reductase, monodehydroascorbate radical reductase, glutathione reductase, and guaiacol peroxidase were affected little or not at all by elevated CO2. When the trees were subjected to drought stress by withholding water, the activities of antioxidative enzymes decreased in leaves of pine and oak grown at ambient CO2 and increased in plants grown at elevated CO2 concentrations. The present results suggest that growth in elevated CO2 might reduce oxidative stress to which leaf tissues are normally exposed and enhance metabolic flexibility to encounter increased stress by increases in antioxidative capacity.

Interspecific variability of δ13C among trees in rainforests of French Guiana: functional groups and canopy integration

Abstract The interspecific variability of sunlit leaf carbon isotope composition (δ13C), an indicator of leaf intrinsic water-use efficiency (WUE, CO2 assimilation rate/leaf conductance for water vapour), was investigated in canopy trees of three lowland rainforest stands in French Guiana, differing in floristic composition and in soil drainage characteristics, but subjected to similar climatic conditions. We sampled leaves with a rifle from 406 trees in total, representing 102 species. Eighteen species were common to the three stands. Mean species δ13C varied over a 6.0‰ range within each stand, corresponding to WUE varying over about a threefold range. Species occurring in at least two stands displayed remarkably stable δ13C values, suggesting a close genetic control of species δ13C. Marked differences in species δ13C values were found with respect to: (1) the leaf phenology pattern (average δ13C=–29.7‰ and –31.0‰ in deciduous-leaved and evergreen-leaved species, respectively), and (2) different types of shade tolerance defined by features reflecting the plasticity of growth dynamics with respect to contrasting light conditions. Heliophilic species exhibited more negative δ13C values (average δ13C=–30.5‰) (i.e. lower WUE) than hemitolerant species (–29.3‰). However, tolerant species (–31.4‰) displayed even more negative δ13C values than heliophilic ones. We could not provide a straightforward ecophysiological interpretation of this result. The negative relationship found between species δ13C and midday leaf water potential (Ψwm) suggests that low δ13C is associated with high whole tree leaf specific hydraulic conductance. Canopy carbon isotope discrimination (ΔA) calculated from the basal area-weighed integral of the species δ13C values was similar in the three stands (average ΔA=23.1‰), despite differences in stand species composition and soil drainage type, reflecting the similar proportions of the three different shade-tolerance types among stands.

Comparison of Quantitative Trait Loci for Adaptive Traits Between Oak and Chestnut Based on an Expressed Sequence Tag Consensus Map

A comparative genetic and QTL mapping was performed between Quercus robur L. and Castanea sativa Mill., two major forest tree species belonging to the Fagaceae family. Oak EST-derived markers (STSs) were used to align the 12 linkage groups of the two species. Fifty-one and 45 STSs were mapped in oak and chestnut, respectively. These STSs, added to SSR markers previously mapped in both species, provided a total number of 55 orthologous molecular markers for comparative mapping within the Fagaceae family. Homeologous genomic regions identified between oak and chestnut allowed us to compare QTL positions for three important adaptive traits. Colocation of the QTL controlling the timing of bud burst was significant between the two species. However, conservation of QTL for height growth was not supported by statistical tests. No QTL for carbon isotope discrimination was conserved between the two species. Putative candidate genes for bud burst can be identified on the basis of colocations between EST-derived markers and QTL.

Functional diversity in an Amazonian rainforest of French Guyana: a dual isotope approach (δ15N and δ13C)

Abstract Functional aspects of biodiversity were investigated in a lowland tropical rainforest in French Guyana (5°2′N, annual precipitation 2200 mm). We assessed leaf δ15N as a presumptive indicator of symbiotic N2 fixation, and leaf and wood cellulose δ13C as an indicator of leaf intrinsic water-use efficiency (CO2 assimilation rate/leaf conductance for water vapour) in dominant trees of 21 species selected for their representativeness in the forest cover, their ecological strategy (pioneers or late successional stage species, shade tolerance) or their potential ability for N2 fixation. Similar measurements were made in trees of native species growing in a nearby plantation after severe perturbation (clear cutting, mechanical soil disturbance). Bulk soil δ15N was spatially quite uniform in the forest (range 3–5‰), whereas average leaf δ15N ranged from −0.3‰ to 3.5‰ in the different species. Three species only, Diplotropis purpurea, Recordoxylon speciosum (Fabaceae), and Sclerolobium melinonii (Caesalpiniaceae), had root bacterial nodules, which was also associated with leaf N concentrations higher than 20 mg g−1. Although nodulated trees displayed significantly lower leaf δ15N values than non-nodulated trees, leaf δ15N did not prove a straightforward indicator of symbiotic fixation, since there was a clear overlap of δ15N values for nodulated and non-nodulated species at the lower end of the δ15N range. Perturbation did not markedly affect the difference δ15Nsoil − δ15Nleaf, and thus the isotopic data provide no evidence of an alteration in the different N acquisition patterns. Extremely large interspecific differences in sunlit leaf δ13C were observed in the forest (average values from −31.4 to −26.7‰), corresponding to intrinsic water-use efficiencies (ratio CO2 assimilation rate/leaf conductance for water vapour) varying over a threefold range. Wood cellulose δ13C was positively related to total leaf δ13C, the former values being 2–3‰ higher than the latter ones. Leaf δ13C was not related to leaf δ15N at either intraspecific or interspecific levels. δ13C of sunlit leaves was highest in shade hemitolerant emergent species and was lower in heliophilic, but also in shade-tolerant species. For a given species, leaf δ13C did not differ between the pristine forest and the disturbed plantation conditions. Our results are not in accord with the concept of existence of functional types of species characterized by common suites of traits underlying niche differentiation; rather, they support the hypothesis that each trait leads to a separate grouping of species.

Pioneer and late stage tropical rainforest tree species (French Guiana) growing under common conditions differ in leaf gas exchange regulation, carbon isotope discrimination and leaf water potential

Leaf gas exchange rates, predawn Ψwp and daily minimum Ψwm leaf water potentials were measured during a wet-to-dry season transition in pioneer (Jacaranda copaia, Goupia glabra andCarapa guianensis) and late stage rainforest tree species (Dicorynia guianensis andEperua falcata) growing in common conditions in artificial stands in French Guiana. Carbon isotope discrimination (Δ) was assessed by measuring the stable carbon isotope composition of the cellulose fraction of wood cores. The Δ values were 2.7‰ higher in the pioneer species than in the late stage species. The calculated time integratedCi values derived from the Δ values averaged 281 μmol mol−1 in the pioneers and 240 μmol mol−1 in the late stage species. The corresponding time-integrated values of intrinsinc water-use efficiency [ratio CO2 assimilation rate (A)/leaf conductance (g)] ranged from 37 to 47 mmol mol−1 in the pioneers and the values were 64 and 74 mmol mol−1 for the two late stage species. The high Δ values were associated—at least inJ. copaia—with high maximumg values and with high plant intrinsinc specific hydraulic conductance [C≔g/(Ψwm−Ψwp], which could reflect a high competitive ability for water and nutrient uptake in the absence of soil drought in the pioneers. A further clear discriminating trait of the pioneer species was the very sensitive stomatal response to drought in the soil, which might be associated with a high vulnerability to cavitation in these species. From a methodological point of view, the results show the relevance of Δ for distinguishing ecophysiological functional types among rainforest trees.

The successional status of tropical rainforest tree species is associated with differences in leaf carbon isotope discrimination and functional traits

We characterised the among species variability in leaf gas exchange and morphological traits under controlled conditions of seedlings of 22 tropical rainforest canopy species to understand the origin of the variability in leaf carbon isotope discrimination (Δ) among species with different growth and dynamic characteristics (successional gradient). Our results first suggest that these species pursue a consistent strategy in terms of Δ throughout their ontogeny (juveniles grown here versus canopy adult trees from the natural forest). Second, leaf Δ was negatively correlated with WUE and N, and positively correlated with gs, but among species differences in Δ were mainly explained by differences in WUE. Finally, species belonging to different successional groups display distinct leaf functional and morphological traits. We confirmed that fast growing early successional species maximise carbon assimilation with high stomatal conductance. In contrast, fast and slow growing late successional species are both characterised by low carbon assimilation values, but by distinct stomatal conductance and leaf morphological features. Along the successional gradient, these differences result in much lower Δ for the intermediate species (i.e. fast growing late successional) as compared to the two other groups.RésuméNous avons caractérisé la variabilité interspécifique des échanges gazeux et des traits morphologiques foliaires en conditions environnementales contrôlées de jeunes plants de 22 espèces d’arbres de la canopée en forêt tropicale humide afin de comprendre l’origine de la variabilité de la discrimination isotopique du carbone foliaire (Δ) observée entre ces espèces présentant des caractéristiques de croissance et de dynamique distinctes (groupes successionnels). Nous montrons premièrement que les espèces tropicales possèdent une stratégie très conservée de Δ au cours de leur ontogénie (juvéniles élevés ici versus arbres adultes de la canopée en forêt naturelle). Deuxièmement, Δ était négativement corrélée à WUE et N, et positivement à gs, mais les différences de Δ entre espèces sont principalement expliquées par des différences de WUE. Enfin, nous montrons que les espèces appartenant à des groupes successionnels distincts présentent des traits fonctionnels et morphologiques foliaires distincts. Nous confirmons que les espèces à croissance rapide qui s’installent en premier au cours de la succession écologique (FE) maximisent A avec de fortes conductances stomatiques. Les espèces climax (qui s’installent en second dans la succession écologique), à croissance rapide (FL) ou à croissance faible (SL), présentent des valeurs de A identiques, mais des valeurs de gs ainsi que des caractéristiques morphologiques foliaires distinctes. Dans la succession écologique, ces différences se traduisent par des valeurs de Δ nettement plus faibles pour les espèces intermédiaires (c’est-à-dire les espèces climax à croissance rapide) par rapport aux deux autres groupes.

Leaf functional response to increasing atmospheric CO2 concentrations over the last century in two northern Amazonian tree species: a historical δ13C and δ18O approach using herbarium samples

We assessed the extent of recent environmental changes on leaf morphological (stomatal density, stomatal surface, leaf mass per unit area) and physiological traits (carbon isotope composition, δ(13)C(leaf) , and discrimination, Δ(13)C(leaf) , oxygen isotope composition, δ(18)O(leaf) ) of two tropical rainforest species (Dicorynia guianensis; Humiria balsamifera) that are abundant in the Guiana shield (Northern Amazonia). Leaf samples were collected in different international herbariums to cover a 200 year time-period (1790-2004) and the whole Guiana shield. Using models describing carbon and oxygen isotope fractionations during photosynthesis, different scenarios of change in intercellular CO(2) concentrations inside the leaf (C(i)), stomatal conductance (g), and photosynthesis (A) were tested in order to understand leaf physiological response to increasing air CO(2) concentrations (C(a)). Our results confirmed that both species displayed physiological response to changing C(a) . For both species, we observed a decrease of about 1.7‰ in δ(13)C(leaf) since 1950, without significant change in Δ(13)C(leaf) and leaf morphological traits. Furthermore, there was no clear change in δ(18)O(leaf) for Humiria over this period. Our simulation approach revealed that an increase in A, rather than a decrease in g, explained the observed trends for these tropical rainforest species, allowing them to maintain a constant ratio of C(i)/C(a) .

Forêts et augmentation de la concentration atmosphérique en CO2 : effets bénéfiques ou adverses ?

Elle affecte directement le fonctionnement des vegetaux et notamment celui des arbres forestiers a travers des effets aboutissant le plus souvent a une augmentation de leur vitesse de croissance. Cet effet a ete mis en evidence pour un grand nombre d’especes forestieres a l’aide d’experimentations basees sur le doublement de la concentration atmospherique en CO2 a partir du niveau actuel (Ceulemans et Mousseau, 1994 ; Saxe et al., 1998). L’effet CO2 est sans doute deja effectif actuellement et constitue un facteur explicatif tres probable dans le determinisme de l’augmentation de la productivite forestiere observee en zone temperee (Spiecker et al., 1996).