Contribution Des Légumineuses, Des Champignons Endophytes Et Mycorhiziens Dans La

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Contribution Des Légumineuses, Des Champignons Endophytes Et Mycorhiziens Dans La Université de Montréal Contribution des légumineuses, des champignons endophytes et mycorhiziens dans la nutrition azotée des prairies indigènes semi-arides Par Rim Klabi Département de sciences biologiques, Centre sur la biodiversité Institut de recherche en biologie végétale Faculté des Arts et des Sciences Thèse présentée à la Faculté des études supérieures en vue de l’obtention du grade de Philosophiæ Doctor (Ph.D.) en sciences biologiques Août, 2015 © Rim Klabi, 2015 i Université de Montréal Faculté des études supérieures et postdoctorales Cette thèse intitulée: Contribution des légumineuses, des champignons endophytes et mycorhiziens dans la nutrition azotée des prairies indigènes semi-arides Présentée par Rim Klabi a été évaluée par un jury composé des personnes suivantes: Luc Brouillet, président-rapporteur Marc St-Arnaud, directeur de recherche Chantal Hamel, co-directrice de recherche Etienne Yergeau, membre du jury Gilles Bélanger, examinateur externe Suzanne Giasson, Représentante du doyen de la FES ii Résumé Les prairies indigènes présentent une source importante d'alimentation pour le pâturage du bétail dans les prairies Canadiennes semi-arides. L'addition de légumineuses fixatrices d'azote et de phosphore dans les prairies indigènes peut améliorer la productivité et la valeur nutritive de fourrage. Ces pratiques peuvent induire des modifications de la structure et de la diversité des communautés fongiques du sol, ce qui peut en retour avoir un impact sur la production et le contenu nutritionnel du fourrage. L’objectif de cette étude était de développer un système de pâturage à bas niveau d’intrants, productif, autonome et durable. À court terme, nous voulions 1) déterminer l'effet des légumineuses (Medicago sativa, une légumineuse cultivée ou Dalea purpurea, une légumineuse indigène) et la fertilité en phosphore du sol sur la productivité et la valeur nutritive des graminées indigènes, comparées avec celles de la graminée introduite Bromus biebersteinii en mélange avec le M. sativa, 2) identifier l'effet de ces pratiques sur la diversité et la structure des communautés des champignons mycorhiziens à arbuscules (CMA) et des champignons totaux, 3) identifier l'effet des légumineuses et des CMA sur les interactions compétitives entre les graminées de saison fraîche et les graminées de saison chaude. Les expériences menées au champ ont montré que M. sativa améliorait les teneurs en azote et en phosphore des graminées indigènes au début de l'été, ainsi que la teneur en azote de la graminée de saison chaude Bouteloua gracilis à la fin de l'été de l'année sèche 2009. Par contre, la fertilité en phosphore du sol n'ait pas affecté la productivité des plantes. D'autre part, l'inclusion des légumineuses augmentait la diversité des CMA dans le mélange de graminées indigènes. Cette modification présentait des corrélations positives avec la productivité et la quantité totale d'azote chez le M. sativa et avec la teneur en phosphore des graminées indigènes, au début de l'été. La structure des communautés de champignons totaux était influencée par l'interaction entre le mélange des espèces et la fertilité en phosphore du sol iii seulement en 2008 (année humide). Cet effet pourrait être lié en partie avec la productivité des plantes et l'humidité du sol. Les expériences menées en chambre de culture ont montré que les CMA peuvent favoriser la productivité des graminées de saison chaude au détriment des graminées de saison fraîche. En effet, Glomus cubense augmentait la productivité de la graminée de saison chaude B. gracilis, en présence de M. sativa. Cet effet pourrait être associé à l’effet négatif du G. cubense sur la fixation de l’azote par le M. sativa et à la diminution de l’efficacité d’utilisation de l'azote de certaines graminées de saison fraîche résultant en une augmentation de la disponibilité de l'azote pour B. gracilis. Par contre, le Glomus sp. augmentait la biomasse de Schizachyrium scoparium, autre graminée de saison chaude, en absence de légumineuse. Ce phénomène pourrait être attribuable à une amélioration de l’efficacité d’utilisation du P de cette graminée. En conclusion, mes travaux de recherche ont montré que la légumineuse cultivée M. sativa peut améliorer la valeur nutritive des graminées indigènes au début de l'été ainsi que celle de la graminée de saison chaude B. gracilis, dans des conditions de sécheresse sévère de la fin de l'été. De plus, l'addition de M. sativa dans le mélange de graminées indigènes peut contribuer à augmenter le nombre des espèces bénéfiques des CMA pour la production et la nutrition du fourrage au début de l'été. Mots clés: prairies semi-arides, graminées indigènes, légumineuses, champignons totaux, champignons mycorhiziens à arbuscule, productivité, nutriments, fourrage. iv Abstract The native grasslands are considered as the main feed source for livestock grazing, in semi-arid regions of the Canadian prairies. The addition of N fixing legumes and phosphorus to semi-arid native grasslands may increase the productivity and nutritive value of forage. However, these practices may also shape the structure and diversity of soil fungal communities which in turn may impact forage production and nutritive value. The global objective of this research was to design productive, self-sustaining, permanent and with low inputs pastures. The specific objectives were 1) to demonstrate the effect of N-fixing legumes (the cultivated legume Medicago sativa or the native legume Dalea purpurea) and soil P fertility on the productivity and nutritive value of native grasses mixes in comparison to the mixture of the introduced grass Bromus biebersteinii and M. sativa, 2) identify the effect of these practices on the diversity and community structure of arbuscular mycorrhizal (AM) fungi and total fungi, and 3) identify the effect of legumes and AM fungi on competitive interactions between native cool-season grasses and native warm- season grasses. The field experiment showed that M. sativa improved the nitrogen and phosphorus concentrations of native grasses mixes early in the summer, as well as the N concentration of the warm-season grass B. gracilis, in late summer of the driest year 2009. In contrast, the soil phosphorus fertility had no effect on plant productivity. On the other hand, the inclusion of legumes to the mix of native grasses generally increased AM fungal diversity. This shift was positively correlated with the productivity and nitrogen uptake by M. sativa and with the phosphorus concentration of native grasses mixes in early summer. The structure of the total fungal community was affected by the interaction between species mixtures and soil P fertility only in the wet year (2008), suggesting that this effect was likely driven in part by plant productivity and soil moisture. v The growth chamber experiment showed that the AM fungi may favoured the growth of warm- season grasses under competition with cool-season grasses. However, Glomus cubense increased the productivity of warm-season grass B. gracilis when growing with M. sativa. This effect might be related to a negative impact of G. cubense on the nitrogen-fixing activity of M. sativa and to a lower N-use efficiency of certain cool-season grasses, which resulted in increased soil N availability for B. gracilis. In contrast, Glomus sp. enhanced the growth of S. scoparium, another warm-season grass in the absence of legumes, and this may be related to improved P-use efficiency in this grass. We concluded that the cultivated legume M. sativa can improve the nutritive value of native grasses mixes early in the summer and also of warm season grass under severe drought conditions in late summer. In addition, the inclusion of M. sativa within native grass mixes may contribute to promote beneficial AM fungi taxa that were involved in forage production and nutrition early in the summer. Keywords: semi-arid grassland, native grasses, legumes, total fungi, AM fungi, productivity, nutrients, forage vi Table des matières Résumé ........................................................................................................................................................ iii Table des matières ..................................................................................................................................... vii Liste des tableaux ........................................................................................................................................ x Liste des figures ......................................................................................................................................... xii Remerciements ......................................................................................................................................... xvi Avant-propos .......................................................................................................................................... xviii Chapitre 1: Introduction générale ............................................................................................................. 1 Chapitre 2: Contribution of legumes to promote the nutritive value of forage in semi-arid grassland pastures ...................................................................................................................................................... 10 2.1 Préface ............................................................................................................................................... 11 2.2 Abstract ............................................................................................................................................
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