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International audience; Coppice forestry is a conventional silvicultural practice that takes advantage of a tree's capacity to respond to disturbances by sprouting. Sprouting capacity is determined by many intrinsic and extrinsic factors such as parent tree age/size, understory light availability and deer browsing, which, under closed canopy conditions are important limiting factors for stump survival or sprout growth. However, the combined effect of potentially confounding abiotic and biotic factors on stump survival and sprout growth remains elusive, even more so under closed canopy conditions. This study aims to quantify the effect of deer browsing on stump mortality and sprout growth under closed-canopy conditions and to compare this effect with other known determinants. Here we show that stump survival and sprout growth in sessile oak (Quercus petraea Matt.) depend on deer browsing, understory light availability and the diameter of the parent tree. By studying paired fenced-unfenced plots, we confirmed that deer browsing decreased stump survival and inhibited sprout growth. Furthermore, by taking advantage of a gradient in understory light availability in monospecific and mixed stands of sessile oak and Scots pine (Pinus sylvestris L.), we showed a clear positive linear relationship between sprout growth and light availability. This relationship explained the observed differences among stand composition types. Finally, we found that increased understory light availability did not compensate for losses due to deer browsing. In the absence of deer browsing, our results demonstrate that sessile oak stumps regenerate well under closed-canopy conditions and maintain a moderate sprouting capacity at least until the age of 70-80 years old. Partial thinning could therefore be a potential tool to renew light-demanding tree species such as sessile oak in mixed high-forest stands. Nevertheless, we do not recommend coppicing sessile oak under closed canopies unless the oak stumps are protected from deer browsing and understory light availability is optimized as much as possible despite closed-canopy-management objectives.

One of the main challenges to predict vegetation dynamics and plant community composition is to identify how biotic and abiotic factors modify the nature and magnitude of plant-plant interactions. The objective of my thesis is to highlight the leading role of multiple interactions (involving more than two biotic compartments) in forest understory dynamics and more specifically regeneration dynamics of target species such as sessile oak. The results emphasize the importance of: (i) effects of deer on response patterns of plant-plant interactions along resource gradients, (ii) indirect interactions, in particular indirect facilitation, (iii) demographic parameters in relation to species phenology, (iv) distinguishing among underlying mechanisms of multiple interactions thanks to environmental monitoring. Thus, my thesis supports the idea that management practices focusing on a single pressure, while ignoring others, are unable to conserve populations of target species because they don’t consider multiple interactions. This highlights the importance to use complementary management strategies to achieve sustainability in the context of global changes (climatic change and deer overabundance).; Un des principaux challenges pour prédire la composition, la structure et la dynamique des communautés végétales est de déterminer comment l’environnement biotique et abiotique va modifier la direction et l’amplitude des interactions entre plantes. L’objectif de ma thèse est de mettre en évidence le rôle prépondérant des interactions complexes (impliquant plus de deux compartiments biotiques) dans la dynamique forestière et plus particulièrement dans la dynamique de régénération des ligneux d’intérêt sylvicole tel que le chêne sessile. Les résultats soulignent l’importance de prendre en compte : (i) l’effet des cervidés sur les patrons de réponses des interactions entre plantes le long des gradients de ressources, (ii) les interactions indirectes et notamment la facilitation indirecte, (iii) des paramètres démographiques différents en relation avec la phénologie des espèces en présence, (iv) la séparation des mécanismes sous-jacents à une interaction écologique multiple via un suivi des conditions environnementales. Ainsi, ma thèse appuie l’idée que les gestions se focalisant sur une unique pression et ignorant les autres pressions ne sont pas capables de maintenir des populations d’espèces cibles car elles ne tiennent pas compte des interactions multiples. Ceci souligne l’importance d’utiliser des stratégies complémentaires pour permettre la pérennité des écosystèmes forestiers et notamment une régénération suffisante dans le cadre des changements globaux que sont le changement climatique et la surabondance de cervidés.

International audience; Mixed plantations have been receiving increasing attention for their documented or supposed potential benefits over monospecific plantations. In particular, the use of neighbouring (or nurse) vegetation around target plants can enhance their performance through limitation of the competing herb layer, and can also improve their morphology. Here we examine the benefits and drawbacks of using neighbour treatments on the response of target trees in open plantations. We set up an experimental plantation in southern France, in which two co-occurring target oak species (the evergreen Quercus ilex and the deciduous Quercus pubescens) were introduced in different neighbour treatments using a tree (Pinus halepensis) and a shrub species (the nitrogen-fixing Coronilla glauca). Oaks were planted with pine neighbours at two densities, with shrub neighbours, in a mixture of pines and shrubs or without neighbours. The ground vegetation was either regularly weeded or left to grow in order to detect any indirect facilitation interactions. Target oak responses (survival, growth, and morphology) were monitored over 7 years. Soil water content and light availability were also measured throughout the experiment. We found competition to be the dominant process driving interactions between neighbours and target tree species. Growth was reduced for both species, but more in weeded than in unweeded treatments, showing an alleviation of competitive interactions by neighbours through limitation of herb layer development. However, in both ground vegetation treatments, growth was severely reduced with Coronilla shrubs. Survival was only significantly impaired for the less shade-tolerant Q. pubescens oak used in combination with shrubs. The negative influence of the neighbour treatments was mainly attributable to light interception, which was particularly high by shrub canopy. Soil moisture was also slightly reduced by shrub neighbours, but it remained high with pines in the unweeded treatment owing to a limited abundance of herbs. However, stem form was improved by the neighbour treatments: oaks developed narrower crowns and greater slenderness with neighbours, whereas oaks in the open showed a bushy morphology. These findings emphasize the need to clearly identify key objectives before implementing mixed plantations (e.g. maximising growth, survival, improving morphology, etc.) and to use active management to control potential undesirable effects of the neighbouring vegetation on target plants.

International audience; Plant structural diversity is usually considered as beneficial for ecosystem functioning. For instance, numerous studies have reported positive species diversity-productivity relationships in plant communities. However, other aspects of structural diversity such as individual size inequality have been far less investigated. In forests, tree size inequality impacts directly tree growth and asymmetric competition, but consequences on forest productivity are still indeterminate. In addition, the effect of tree size inequality on productivity is likely to vary with species shade-tolerance, a key ecological characteristic controlling asymmetric competition and light resource acquisition. Using plot data from the French National Geographic Agency, we studied the response of stand productivity to size inequality for ten forest species differing in shade tolerance. We fitted a basal area stand production model that included abiotic factors, stand density, stand development stage and a tree size inequality index. Then, using a forest dynamics model we explored whether mechanisms of light interception and light use efficiency could explain the tree size inequality effect observed for three of the ten species studied. Size inequality negatively affected basal area increment for seven out of the ten species investigated. However, this effect was not related to the shade tolerance of these species. According to the model simulations, the negative tree size inequality effect could result both from reduced total stand light interception and reduced light use efficiency. Our results demonstrate that negative relationships between size inequality and productivity may be the rule in tree populations. The lack of effect of shade tolerance indicates compensatory mechanisms between effect on light availability and response to light availability. Such a pattern deserves further investigations for mixed forestswhere complementarity effects between species are involved. When studying the effect of structural diversity on ecosystem productivity, tree size inequality is a major facet that should be taken into account.

[Departement_IRSTEA]Territoires [TR1_IRSTEA]SEDYVIN; International audience; Mixed plantations have been receiving increasing attention for their documented or supposed potential benefits over monospecific plantations. In particular, the use of neighbouring (or nurse) vegetation around target plants can enhance their performance through limitation of the competing herb layer, and can also improve their morphology. Here we examine the benefits and drawbacks of using neighbour treatments on the response of target trees in open plantations. We set up an experimental plantation in southern France, in which two co-occurring target oak species (the evergreen Quercus ilex and the deciduous Quercus pubescens) were introduced in different neighbour treatments using a tree (Pinus halepensis) and a shrub species (the nitrogen-fixing Coronilla glauca). Oaks were planted with pine neighbours at two densities, with shrub neighbours, in a mixture of pines and shrubs or without neighbours. The ground vegetation was either regularly weeded or left to grow in order to detect any indirect facilitation interactions. Target oak responses (survival, growth, and morphology) were monitored over 7 years. Soil water content and light availability were also measured throughout the experiment. We found competition to be the dominant process driving interactions between neighbours and target tree species. Growth was reduced for both species, but more in weeded than in unweeded treatments, showing an alleviation of competitive interactions by neighbours through limitation of herb layer development. However, in both ground vegetation treatments, growth was severely reduced with Coronilla shrubs. Survival was only significantly impaired for the less shade-tolerant Q. pubescens oak used in combination with shrubs. The negative influence of the neighbour treatments was mainly attributable to light interception, which was particularly high by shrub canopy. Soil moisture was also slightly reduced by shrub neighbours, but it remained high with pines in the unweeded treatment owing to a limited abundance of herbs. However, stem form was improved by the neighbour treatments: oaks developed narrower crowns and greater slenderness with neighbours, whereas oaks in the open showed a bushy morphology. These findings emphasize the need to clearly identify key objectives before implementing mixed plantations (e.g. maximising growth, survival, improving morphology, etc.) and to use active management to control potential undesirable effects of the neighbouring vegetation on target plants.

/ Identifier les facteurs qui influencent le succès des plantes exotiques envahissantes est un enjeu majeur pour leur contrôle. Les méthodes classiques de gestion des plantes exotiques envahissantes montrent leurs limites et l'utilisation des propriétés compétitives des communautés végétales suscite de plus en plus d'intérêt. Toutefois si la résistance biotique des communautés envahies, et en particulier la compétition interspécifique fait partie des mécanismes impliqués dans la régulation des espèces exotiques envahissantes, son importance relative est mal connue. Dans cette étude, nous avons cherché à identifier les facteurs, biotiques et abiotiques, qui influencent la performance végétative de renouées envahissantes établies (Fallopia sp.). Les renouées asiatiques se développent préférentiellement dans les habitats ouverts perturbés, ce qui suggère un rôle prépondérant de la compétition pour la lumière. Nous avons testé cette hypothèse par des observations sur des taches de renouées établies dans les Alpes. Nous avons d'abord mesuré leur performance (production végétative) dans des conditions abiotiques variées. Puis, à partir de traits biologiques liés à la compétition pour la lumière, nous avons caractérisé la structure fonctionnelle des communautés végétales voisines au travers de deux indices complémentaires : la moyenne pondérée de la communauté (community weighted mean) et l'indice de Rao. Les résultats montrent que les renouées envahissantes sont moins vigoureuses en condition de faible lumière tandis que les autres facteurs abiotiques n'influencent pas leur performance. La disponibilité en lumière dépend fortement de la structure fonctionnelle des communautés végétales voisines en termes de hauteur et en particulier de la moyenne pondérée des hauteurs de la communauté. Cette étude confirme l'importance de la compétition pour la lumière sur le développement végétatif des renouées et offre des perspectives en termes de restauration des communautés envahies.

International audience; Context: Developing silvicultural methods to help Mediterranean forests adapt to climate change is of high importance. Introducing resprouting hardwood species below pine stands is expected to promote diversity and resilience of these stands, particularly to forest fires. Aims: To examine how the intensity of pine thinning influences understory micro environment and the establishment of various hardwood seedlings in two Mediterranean sites. Methods: Aleppo pine stands were thinned down to three levels of basal area (uncut: 30 m²/ha, moderate thinning: 13-20 m²/ha, heavy thinning: 7-10 m²/ha) at two Mediterranean sites (South-East France and South-East Spain). Seedlings of six hardwood species were introduced in the understory and their survival and growth were monitored and related to changes in micro-environment induced by thinning. Results: At both sites, thinning improved light availability and seedling diameter increment of all target species. Thinning increased extreme temperature and evaporative demand. Heavy thinning increased summer soil moisture in SE Spain but not in SE France. The worst conditions for seedling survival were reached under uncut stands in SE France and low-density stands in SE Spain. Conclusion: Thinning in pine stands accelerated seedling growth but excessive thinning worsened summer drought and affected seedling survival. Moderate thinning (15-20 m²/ha) seems to be the best option in support of the introduction of hardwoods in the understory, which can improve forest diversity and resilience in the future.

Les travaux présentés dans cet article résultent d’une thèse qui s’inscrit dans la convention de recherche Cemagref-ONF et s’intéresse au renouvellement des peuplements mélangés chêne – pin en forêt d’Orléans. Vu la différence d’exploitabilité des deux essences cela suppose de doser le couvert arboré pour permettre l’installation de semis sans les exposer à une trop forte concurrence de la végétation. Il s’agit donc de préciser les besoins en lumière des semis de pin sylvestre et d’étudier les mécanismes de compétition entre semis et végétation interférente.

Calluna vulgaris , Molinia caerulea and Pteridium aquilinum are three forest understorey species that compete with tree seedlings for environmental resources. Forest managers therefore seek to control their growth, which is driven in part by the light available in the forest understorey. An experiment was set up in a nursery in which the three understorey species C. vulgaris , M. caerulea and P. aquilinum were planted at six light levels (6, 11, 22, 36, 48 and 100% of full sunlight) replicated in three blocks. The different light levels other than 100% were obtained using neutral shading nets. The response of the three species to light availability was estimated after two growing seasons by their cover and height, their morphology, assessed by the foliar tuft height-to-width ratio for C. vulgaris and M. caerulea and by the frond height-to-length ratio for P. aquilinum , and their functional acclimation, assessed by a leaf trait (leaf mass on an area basis, LMA). Vegetation cover changed with light availability following a bell-shaped curve. By contrast, vegetation height remained approximately the same, irrespective of the light treatment, except for P. aquilinum , which grew higher than the other two species at medium and high light levels. As a consequence, vegetation individuals were more spindly in a dark environment. From the lowest light treatment, P. aquilinum cover was high (75%) and intercepted a larger proportion of the light than the other two species. This species thus proved the most competitive for light. All three species showed a notable acclimation to light availability by their LMA, though in different ways. The LMA of C. vulgaris decreased in shaded conditions by an increase in leaf area, whereas decrease in LMA for P. aquilinum was due to a decrease in leaf mass. For M. caerulea , the LMA remained stable whatever the light conditions by a simultaneous increase in both leaf mass and leaf area in shaded conditions. Consequently, the in-forest development of C. vulgaris and M. caerulea could be managed through small adult canopy thinnings, as both species grew well from light levels above 20%. Conversely, it seems unrealistic to manage P. aquilinum in this way as its development was almost maximal even at very low light levels.

Diplôme : IUT

International audience; Pinus sylvestris is a pioneer species and as such is relatively light-demanding. Therefore, its natural regeneration may be inhibited by some forest understorey species that develop with light and can then reduce light as well as soil resources (i.e. nutrients and water) availability for the pine seedlings. To better quantify these effects, we designed two experiments in a nursery. The first one aimed at studying the influence of density of three common understorey species in temperate forests (Calluna vulgaris, Pteridium aquilinum and Molinia caerulea) on pine seedlings survival and growth. The purpose of the second experiment was to quantify the development of these three species relatively to light that varied in our experiment from 6 to 100% of relative light. An increase in weed density, i.e. a concomitant decrease of light availability and soil moisture content, resulted in reduced pine seedlings growth (diameter and biomass) and higher stem height/diameter ratio but had no significant effect on seedlings survival. Height and cover of the three weedy species followed a non linear relationship relative to light availability. Results are discussed according to the quantity of available light as well as its quality assessed through the red/far red ratio, both for the pine seedlings development and weed species height and cover, and their interactions.

International audience; Pinus pinea is one of the most valuable species used in Tunisia in the reforestation program. This species is shade-intolerant and hence needs light to correctly regenerate. It is also influenced by the understorey vegetation and the litter biomass, both also correlated to light availability. To quantify the importance of these different factors, the natural regeneration and evolution of the biomass of understorey vegetation and litter were studied in artificial forests of Pinus pinea located in the coastal dunes of Nefza-Tabarka, and thinned to reach different levels of light. Overstorey cover was measured on 37 plots (500 m2 each) and the biomass of the different species of the understorey and the needle litter was recorded in 8 subplots (0,25m2). The number and age of Pinus pinea seedlings were also determined in each plot. Results showed that the biomass of the understorey vegetation, as well as the number of pine seedlings increased with light availability and concomitantly a decrease of the litter biomass. We will discuss the interactions between the different factors and give practical recommendations to better manage pine stands in the area.

Les hêtraies mélangées sont un enjeu important pour la forêt française. De nouveaux outils existent ou sont en cours de développement, pour mettre en ½uvre une gestion multifonctionnelle de ces forêts, basés sur une modélisation de la dynamique du mélange et des différentes essences qui les constituent. / Mixed beech stands are important for the French forest. New tools are developed for a better multi-functional management of these forests, based on the modelling of their dynamics.

Links between architectural stages of development (ASD), photosynthetic capacities and leaf characteristics in Tachigali melinonii (Caesalpiniaceae) were investigated in the tropical forest of French Guiana. Three ASD were sampled under three light conditions to elucidate the impact of ASD- and light-related changes on photosynthetic functional traits. Photosynthetic parameters (maximum rate in RubisCO carboxylation (Vcmax) and maximum rate of electron transport (Jmax) were estimated by adjusting a biochemical model (Farquhar et al., 1980) to response curves of CO2 assimilation rate to intercellular CO2 concentration, under saturated light. In addition, N and C foliar concentrations were analysed and N allocations to the carboxylation and bioenergetic processus of photosynthesis were calculated using the model of Niinemets and Tenhunen (1997). The variations in ecophysiological parameters observed were due to leaf anatomical and structural changes instead of changes in photosynthetic components, suggesting adjustments to light conditions. Tachigali melinonii thus appears to display morphological and anatomical plasticity at different ASDs and under different light conditions., Une espèce forestière de Guyane Française, Tachigali melinonii (Caesalipiniaceae), a été choisie afin de regarder la variation des traits fonctionnels relatifs à la photosynthèse en fonction de la disponibilité en lumière et en fonction de son stade de développement architectural. Du fait du tempérament de l'espèce vis-à-vis de la lumière de type «struggling gambler» (ombrage nécessaire aux jeunes stades puis développement en pleine lumière), trois stades clefs de développement parmi la séquence de différenciation de l'espèce ainsi que trois environnements lumineux ont été choisis. Les capacités photosynthétiques foliaires ont été estimées à partir de l'ajustement de paramètres du modèle biochimique de photosynthèse foliaire (Farquhar et al. 1980) sur des courbes de réponses de l'assimilation de CO² (A) à des concentrations intracellulaires en CO² (Ci) croissantes, en condition de lumière saturante. La vitesse maximale de carboxylation Vcmax et le flux maximal d'électrons photosynthétiques Jmax ont ainsi été estimés et ensuite reliés aux caractéristiques structurales foliaires, notamment la concentration en azote, en carbone ainsi qu'avec la densité et l’épaisseur. Les fractions d'azote allouées aux processus photosynthétiques de carboxylation et de bioénergétique ont été calculées. Des variations, dues à un effet du stade de développement sur les paramètres étudiés ainsi qu’à un effet de la quantité de lumière disponible, ont été constatées. Ces variations sont surtout liées à des ajustements anatomiques et structuraux foliaires et non à des modifications de l’appareil photosynthétique. Tachigali melinonii semble donc modifier ses caractéristiques structurales foliaires en fonction de son stade de développement, ce qui suppose un contrôle ontogénique partiel, et en fonction de la disponibilité en lumière.