<p>In this thesis, I experimentally investigate feeding selectivity in fallow deer (Dama dama), with respect to plant secondary compounds, especially tannins, which can decrease the quality of foods. I found that fallow deer avoided foods with higher amounts of tannic acid and Quebracho tannin, even though the deer ate some high-tannin food. The food choice was strongly dependent on the context in which the food was presented, so that the food choice in relation to tannin content was relative rather than absolute. When high-tannin food occurred at low frequency, the deer ate proportionally less from this type of food, at least when the difference in tannin content between the two foods was large. A basic implication is that an unpalatable plant type could benefit from its unpalatability, especially when occurring at low frequency. In experiments with two patches, the finding of a stronger within- than between-patch selectivity was mirrored in associational effects. First, low-tannin, palatable food was more eaten when occurring in a high-tannin patch, which corresponds to neighbour contrast susceptibility. Second, high-tannin, unpalatable food in a less defended patch was less eaten, which corresponds to neighbour contrast defence. A proximate cause of the associational effects can be the presence of a simultaneous negative contrast, which was experimentally demonstrated in an additional study. Individual differences in selectivity were present early in life and were consistent over five years, and selectivity was correlated with foraging exploratory behaviour. The results from this thesis suggest that fallow deer are selective in their food choice with respect to tannins from the beginning, and that the frequency of occurrence of different foods, but also the distance between foods and the complexity of presentation, influence the food choice. It is also suggested that a foraging behavioural syndrome is present in mammalian herbivores.</p>

In this thesis, I experimentally investigate feeding selectivity in fallow deer (Dama dama), with respect to plant secondary compounds, especially tannins, which can decrease the quality of foods. I found that fallow deer avoided foods with higher amounts of tannic acid and Quebracho tannin, even though the deer ate some high-tannin food. The food choice was strongly dependent on the context in which the food was presented, so that the food choice in relation to tannin content was relative rather than absolute. When high-tannin food occurred at low frequency, the deer ate proportionally less from this type of food, at least when the difference in tannin content between the two foods was large. A basic implication is that an unpalatable plant type could benefit from its unpalatability, especially when occurring at low frequency. In experiments with two patches, the finding of a stronger within- than between-patch selectivity was mirrored in associational effects. First, low-tannin, palatable food was more eaten when occurring in a high-tannin patch, which corresponds to neighbour contrast susceptibility. Second, high-tannin, unpalatable food in a less defended patch was less eaten, which corresponds to neighbour contrast defence. A proximate cause of the associational effects can be the presence of a simultaneous negative contrast, which was experimentally demonstrated in an additional study. Individual differences in selectivity were present early in life and were consistent over five years, and selectivity was correlated with foraging exploratory behaviour. The results from this thesis suggest that fallow deer are selective in their food choice with respect to tannins from the beginning, and that the frequency of occurrence of different foods, but also the distance between foods and the complexity of presentation, influence the food choice. It is also suggested that a foraging behavioural syndrome is present in mammalian herbivores.

Les légumineuses permettent de réduire les intrants en élevage (engrais, concentrés) en raison notamment de leurs niveaux élevés en protéines. Cependant, à la fois pendant le processus d'ensilage et celui de fermentation dans le rumen, les protéines peuvent subir une importante dégradation, ce qui affecte la valeur nutritive des fourrages et induit des rejets d'azote (N) importants, notamment dans l'urine. Certaines légumineuses peuvent alors être d'un intérêt particulier car elles produisent des composés secondaires qui peuvent modifier positivement les processus fermentaires et digestifs. Ainsi, les tannins condensés (CT) présents dans le sainfoin (SF, Onobrychis viciifolia) sont capables de se lier aux protéines, réduisant leur dégradation dans le silo et le rumen et se traduisant par un transfert de l'excrétion d'azote de l'urine vers les fèces. Le trèfle violet (RC, Trifolium pratense) contient la polyphénoloxydase (PPO), une enzyme qui catalyse l'oxydation de différents composés phénoliques en quinones. Comme les CTs, les quinones sont capables de former des complexes avec les protéines permettant de réduire leur dégradation dans le silo et le rumen. L'objectif de cette thèse était alors d'étudier et de quantifier les bénéfices potentiels de l'utilisation de ces deux espèces de légumineuses bioactives sur i) la qualité et la conservation des ensilages, ii) la fermentation ruminale, l'efficacité digestive et les performances des ovins, et iii) l'empreinte environnementale (excrétion d'N et de CH4). Nous avons effectué deux essais in vitro et deux essais in vivo, basés sur des ensilages composés de ces deux légumineuses, seules ou en mélange avec une graminée (la fléole- T, Phleum pratense L.) qui nous a servie de contrôle. Les essais in vitro nous ont permis de nous focaliser sur la qualité et la conservation des ensilages ainsi que sur la fermentation ruminale, tandis que les essais in vivo se sont concentrés sur la performance et l'efficacité digestive des agneaux, ainsi que sur leur bilan azoté et leurs émissions de CH4. L'inclusion de légumineuses bioactives dans les ensilages d'herbe a amélioré la qualité du fourrage, la fermentation pendant le processus d'ensilage ainsi que la protection des protéines contre une dégradation au sein du silo et du rumen. Globalement, l'alimentation des agneaux avec des mélanges comportant ces légumineuses s'est traduite par une augmentation de l'ingestion de matière sèche, en comparaison des agneaux alimentés avec la graminée pure. Néanmoins, en raison de la digestibilité nettement plus faible de T-SF, probablement due à une composition et une nature des fibres différentes ainsi qu'à la présence de CT, les agneaux ayant reçu T-SF ont montré une ingestion et des performances plus faibles que ceux ayant reçu les ensilages contenant RC. Dans le rumen, il semble que les protéines de RC aient été plus protégées de la dégradation que celles de SF, tandis que dans la suite du tractus digestif, les complexes formés entre protéines et CT (avec SF) se seraient moins dissociés que ceux formés entre protéines et quinones (avec RC), ce qui pourrait en partie expliquer le transfert d'excrétion de l'N de l'urine vers les fèces, observé chez les agneaux alimentés avec T-SF et bénéfique pour l'environnement. SF a également permis de réduire légèrement les émissions de CH4. Ainsi, utiliser des légumineuses bioactives dans les pratiques d'alimentation des ruminants apparaît une stratégie prometteuse pour fournir des produits animaux de façon plus durable. Nos résultats montrent que chaque espèce apporte des avantages différents, plutôt orientés vers la qualité de l'aliment et les performances animales pour RC mais plutôt orientés vers la réduction des rejets pour SF. Des recherches complémentaires sont donc nécessaires pour mieux caractériser ces avantages et élargir les investigations à d'autres espèces, d'autres mélanges et d'autres bénéfices potentiels. (...) / Fodder legume species allow to reduce inputs in livestock breeding systems (fertilizer, concentrates) notably because they contain high levels of crude proteins which are of primary importance in ruminant nutrition. However, during both silage and rumen fermentation processes, proteins are submitted to degradation which affects forage nutritive value and leads to nitrogen (N) losses notably via urine. Some specific legumes can then be of particular interest as they produce plant secondary compounds that can positively affect silage and digestive processes. Condensed tannins (CTs) present in sainfoin (SF, Onobrychis viciifolia) are able to bind with proteins thereby reducing their degradation in the silo and the rumen, resulting in a shift in N excretion from urine to faeces. Red clover (RC, Trifolium pratense) contains polyphenol oxidase (PPO), an enzyme that catalyses the oxidation of different phenolics into quinones. As CTs, quinones are able to form complexes with proteins that will similarly reduce their degradation in the silo and the rumen. The aim of this thesis was to investigate and quantify the potential benefits of using these two bioactive legume species on i) quality and conservation of silages, ii) rumen fermentation, digestive efficiency and sheep performance, and iii) environmental footprint (N excretion and CH4 emissions). We conducted two in vitro and two in vivo trials which were based on silages of pure legumes or of different mixtures with the grass species (timothy T, Phleum pratense L.), which served as control. In the in vitro trials, we focussed on silage quality, silage conservation and rumen fermentation, while in the in vivo trials, we focussed on lambs' performance, digestion efficiency, N balance and CH4 emissions. Including bioactive legumes in mixtures with grass improved, compared to pure grass, forage quality and fermentation during the silage making process, as well as proteins' protection from degradation within both the silos and the rumen. Lambs fed with the mixtures involving legumes responded with an increase in DM intake compared to their counterparts fed with T. Nevertheless, due to a possibly different fibre composition and to the presence of CT which impaired SF digestibility, lambs that consumed T-SF showed lower intake and performance than those that received RC-containing silages. In the rumen, RC proteins appeared more protected from degradation than SF ones, while in the subsequent parts of the digestive tract, the proteins-CT complexes (from SF) might less dissociate than the proteins-quinones ones (from RC). This could partly explain the environment-friendly shift in N excretion from urine to faeces when animals are fed with T-SF. SF also allowed to slightly reduce CH4 emissions. Thus, utilizing bioactive legumes in livestock feeding practices is a promising strategy to produce animal products more sustainably. Our results show different benefits relative to the bioactive legume species involved, directed towards boosted forage quality and animals' performance for RC but towards lowered wastes for SF. Further research is thus needed to better characterize these benefits and enlarging investigations to other plant species, mixtures and potential benefits (e.g. health). This will help to determine the appropriate choice of plant species according to the objectives.