965 resultados para Nitrogen uptake kinetics
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The effect of molecular nitrogen exposure on the InP(100) surface modified by the alkali metal K overlayer is investigated by core-level photoemission spectroscopy using synchrotron radiation. The alkali metal covered surface exhibits reasonable nitrogen uptake at room temperature, and results in the formation of a P3N5 nitride complex. Flash annealing at 400 degrees C greatly enhanced the formation of this kind of nitride complex. Above 500 degrees C, the nitride complex dissolved completely. (C) 1997 American Vacuum Society.
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以采自于黄土高原差异较大的25个农田耕层石灰性土壤为供试土样,以淋洗和未淋洗土壤起始NO3--N小麦和玉米两季盆栽试验作物累积吸氮量为参比,对8种反映旱地土壤供氮能力的化学方法进行比较研究。结果表明,在一定程度上,可用有密切关系的土壤全氮或有机质,反映有机氮或全氮存在较大差异的土壤供氮能力,但其灵敏性较差。石灰性土壤矿质氮,特别是NO3--N与未淋洗土壤起始NO3--N作物吸氮量之间有较高相关性(r=0.884,P<0.01),而与淋洗土壤起始NO3--N作物吸氮量间相关系数仅为0.472(P<0.05),说明矿质氮可反映土壤当前供氮水平,而不能反映土壤潜在供氮能力;石灰性土壤起始NO3--N对各化学方法与作物吸氮量之间相关性影响较大。酸性高锰酸钾法既可反映土壤潜在供氮能力,也可反映土壤总供氮能力;酸性高锰酸钾法的修订方法,即硫酸—高锰酸钾法提取出的NH4+-N值接近于KCl水浴法提取出的NH4+-N值,该方法在反映土壤总供氮能力方面与酸性高锰酸钾法相当,但在反映土壤潜在供氮能力方面不及酸性高锰酸钾法优越。KCl水浴法在评价石灰性土壤供氮能力方面,与酸性高锰酸钾法的效果基本相同;沸水浸取法和NaHCO3-UV...
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本文采用批次培养和半连续培养的方法,研究了中国近海的主要赤潮生物之一尖刺菱形藻的营养盐吸收动力学和营养盐限制下的生长进行了研究。1)中国近海的尖刺菱形藻为尖刺型,不能形成软骨藻酸;2)尖刺菱形藻是一种典型的广温广盐的近岸型海洋浮游硅藻,其适应不同环境的能力较强;3)尖刺菱形藻的营养盐吸收动力学符合米氏酶动力学方程V=V_mXS/(Ks+S);4)在稳定状态下,米氏方程与Droop方程一起,与Mond方程一样,都可以很好地描述尖刺菱形藻的生长率与外界环境中限制性营养盐浓度之间的关系 ;5)与其他浮游植物相比较,尖刺菱形藻吸收无机氮的能力较强,而吸收无机磷的能力相对较弱。
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There is excess nitrate (NO3) in the Pearl River coastal plume in the southern waters of Hong Kong in summer. We hypothesize that phosphorus (P) limitation controls the utilization of excess NO3 due to the high N:P ratio in the Pearl River. To test this hypothesis, we conducted two 1-day cruises on July 13 and 19, 2000 to examine the response of the phytoplankton to P additions with respect to changes in biomass, uptake of nutrients and nutrient uptake ratios using a batch incubation of natural water samples collected from the Pearl River estuary and adjacent coastal waters. At a station (E1, salinity =5) in the Pearl River estuary, the N/P ratio at the surface was 46:1, (64 muM DIN: 1.3 muM PO4) and decreased to 24:1 (12 muM DIN: 0.5 muM PO4) downstream at a station (Stn 26, salinity =26) in the coastal plume south of Hong Kong. Without a P addition, NO3 in the water samples collected at E1 could not be depleted during a 9 day incubation (similar to20 muM NO3 remaining). With a P addition, NO3 disappeared completely on day 6 with the depletion of the added PO4 (2-3 muM). This was also true for a station, E4 (salinity= 15) further downstream, but within the estuary. At Stn 26, in the coastal plume south of Hong Kong, NO3 (similar to11.5 muM) was eventually depleted without the addition of PO4, but it took 8 days instead of 5 days for Stn E4. The uptake ratio of dissolved inorganic nitrogen (DIN) to PO4, without a P addition was 51:1, 43:1 and 46:1 for Stns E1, E4 and 26, respectively. With a P addition, the DIN/PO4 uptake ratio decreased to 20:1, 14:1 and 12:1, respectively, for the 3 stations. These results clearly indicate potential P limitation to utilization of NO3 in the Pearl River estuary, resulting in excess NO3 in waters of the coastal plume downstream of the estuary, some of which would eventually be transported offshore. High uptake ratios of N:P without a P addition (43N:1P) suggest that phytoplankton have a nitrogen uptake capacity in excess of the Redfield ratio of 16N: 1P by 2.5-3 times. The value of 2.5-3 times was likely a maximum that should have contained a contribution of P released from desorption of P from sediments or from regeneration by zooplankton grazing and bacterial activity during the incubation of natural water samples. Without a P addition, however, phytoplankton biomass did not increase. This means that P turnover rates or regeneration may allow phytoplankton to take up additional N in excess of the Redfield ratio and store it, but without increasing the algal biomass. Therefore, high ambient N:P ratios in excess of the Redfield ratio do indicate potential P limitation to phytoplankton biomass in this estuarine coastal ecosystem. (C) 2004 Elsevier Ltd. All rights reserved.
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Seasonal investigations of size-fractionated biomass and production were carried out from February 1992 to May 1993 in Jiaozhou Bay, China. Microplankton assemblages were separated into three fractions: pico-(0.7-2 mu m), nano- (2-20 mu m) and netplankton (20-200 mu m). The biomass was measured as chlorophyll a (Chl a), particulate organic carbon (POC) and particulate organic nitrogen (PON). The production was determined by C-14 and N-15 tracer techniques. The seasonal patterns in biomass, though variable, were characterized by higher values in spring and lower values in autumn and summer (for Chl a only). The seasonal patterns in production, on the other hand, were more clear with higher values occurring in summer and spring, and lower values occurring in autumn and winter. Averaged over the whole study period, the respective proportions of total biomass accounted for by net-, nano- and picoplankton were 26, 45 and 29% for Chl a, 32, 33 and 35% for POC, and 26, 32 and 42% for PON. The contributions to total primary production by net-, nano- and picoplankton were 31, 35 and 34%, respectively. The respective proportions of total NH4+-N uptake accounted for by net-, nano- and picoplankton were 28, 33 and 39% in the daytime, and 10, 29 and 61% at night. The respective contributions to total NO3--N uptake by net-, nano- and picoplankton were 37, 40 and 23% in the daytime, and 13, 23 and 64% at night. Some comprehensive ratios, including C/N biomass ratio, Chl a/C ratio, C uptake/Chl a ratio, C:N uptake ratio and the f-ratio, were also calculated size separately, and their biological and ecological meanings are discussed.
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Visando a elucidar alguns aspectos do metabolismo do nitrogenio por plantas de Panicum maximum, estudaram-se os efeitos da proporcao NH +4/NO-3 na variacao do pH das solucoes nutritivas de tratamento, no acumulo de materia seca e na distribuicaoo de nitrogenio na planta. Para tal, as plantas foram cultivadas em camara de crescimento na solucao nutritiva (pH 5,5) completa, contendo nitrogenio somente na forma nitrica. Aos 25 dias depois da germinacao, as plantas foram submetidas a 100 ppm de nitrogenio fornecido nas formas nitrica e amoniacal ou amonical, nas seguintes proporcoes 0/100,25/75,50/50,75/25 e 100/0, durante quatro dias. O pH das solucoes nutritivas foi medido e corrigido diariamente. Os resultados demostraram que o pH aumentou na presenca exclusiva de nitrato (pH 6,5) e decresceu na presenca exclusiva de amonio (pH 3,5). Na presenca de ambas as formas de nitrogenio, o pH permaneceu em torno de 5,0 nos dois primeiros dias entao diminui para cerca de 4,5 nos dias subsequentes. Esses resultados indicam que as plantas de Panicum maximum absorveram o nitrato, inicialmente, porem com o tempo adaptaram-se a presenca de amonio e tenderam a absorver ambas as formas. As plantas que receberam nitrato, combinados ou não com amonio, apresentaram o peso da materia seca, N-total e N-soluvel total superiores as que receberam somente amonio como fonte de N, havenso pequena tendencia de maior acumulo na presenca de ambas as fontes. No tratamento 100% de amonio, pN-total e o N-soluvel total diminuiram apreciavelmente, indicando baixa capacidade de essa especie utilizar amonio como fonte única de nitrogenio. Os resultados indicam que, embora as plantas possa absorver as duas formas de nitrogenio, a combinacao entre ambas parece ser a forma ideal para o crescimento das mesmas.
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Intercellular adhesion molecule 1 (ICAM-1) is a transmembrane protein found on the surface of vascular endothelial cells (ECs). Its expression is upregulated at inflammatory sites, allowing for targeted delivery of therapeutics using ICAM-1-binding drug carriers. Engagement of multiple copies of ICAM-1 by these drug carriers induces cell adhesion molecule (CAM)-mediated endocytosis, which results in trafficking of carriers to lysosomes and across ECs. Knowledge about the regulation behind CAM-mediated endocytosis can help improve drug delivery, but questions remain about these regulatory mechanisms. Furthermore, little is known about the natural function of this endocytic pathway. To address these gaps in knowledge, we focused on two natural binding partners of ICAM-1 that potentially elicit CAM-mediated endocytosis: leukocytes (which bind ICAM-1 via β2 integrins) and fibrin polymers (a main component of blood clots which binds ICAM-1 via the γ3 sequence). First, inspired by properties of these natural binding partners, we varied the size and targeting moiety of model drug carriers to determine how these parameters affect CAM-mediated endocytosis. Increasing ICAM-1-targeted carrier size slowed carrier uptake kinetics, reduced carrier trafficking to lysosomes, and increased carrier transport across ECs. Changing targeting moieties from antibodies to peptides decreased particle binding and uptake, lowered trafficking to lysosomes, and increased transport across ECs. Second, using cell culture models of leukocyte/EC interactions, inhibiting regulatory elements of the CAM-mediated pathway disrupted leukocyte sampling, a process crucial to leukocyte crossing of endothelial layers (transmigration). This inhibition also decreased leukocyte transmigration across ECs, specifically through the transcellular route, which occurs through a single EC without disassembly of cell-cell junctions. Third, fibrin meshes, which mimic blood clot fragments/remnants, bound to ECs at ICAM-1-enriched sites and were internalized by the endothelium. Inhibiting the CAM-mediated pathway disrupted this uptake. Following endocytosis, fibrin meshes trafficked to lysosomes where they were degraded. In mouse models, CAM-mediated endocytosis of fibrin meshes appeared to remove fibrin remnants at the endothelial surface, preventing re-initiation of the coagulation cascade. Overall, these results support a link between CAM-mediated endocytosis and leukocyte transmigration as well as uptake of fibrin materials by ECs. Furthermore, these results will guide the future design of ICAM-1-targeted carrier-assisted therapies.
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Arsenate resistance is exhibited by the ericoid mycorrhizal fungus Hymenoscyphus ericae collected from As-contaminated mine soils. To investigate the mechanism of arsenate resistance, uptake kinetics for arsenate (H(2)AsO(4)(-)), arsenite (H(3)AsO(3)), and phosphate (H(2)PO(4)(-)) were determined in both arsenate-resistant and -non-resistant H. ericae. The uptake kinetics of H(2)AsO(4)(-), H(3)AsO(3), and H(2)PO(4)(-) in both resistant and non-resistant isolates were similar. The presence of 5.0 microM H(2)PO(4)(-) repressed uptake of H(2)AsO(4)(-) and exposure to 0.75 mM H(2)AsO(4)(-) repressed H(2)PO(4)(-) uptake in both H. ericae. Mine site H. ericae demonstrated an enhanced As efflux mechanism in comparison with non-resistant H. ericae and lost approximately 90% of preloaded cellular As (1-h uptake of 0.22 micromol g(-1) dry weight h(-1) H(2)AsO(4)(-)) over a 5-h period in comparison with non-resistant H. ericae, which lost 40% of their total absorbed H(2)AsO(4)(-). As lost from the fungal tissue was in the form of H(3)AsO(3). The results of the present study demonstrate an enhanced H(3)AsO(3) efflux system operating in mine site H. ericae as a mechanism for H(2)AsO(4)(-) resistance. The ecological significance of this mechanism of arsenate resistance is discussed.
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Significant genotypic difference in response to arsenate toxicity in rice (Oryza sativa) was investigated in root elongation, arsenate uptake kinetics, physiological and biochemical response and arsenic (As) speciation. Uptake kinetics data showed that P-deprived genotype 94D-54 had a little higher As uptake than P-deprived 94D-64, but the difference was not large enough to cause acute toxicity in P-deprived 94D-54. There was no difference in tissue P concentrations between the two genotypes under P deficient conditions. In addition, arsenic speciation in plant tissues (using high performance liquid chromatography-inductively coupled plasma mass spectrometry) was not different between P pretreatments and between genotypes. P-deprived genotype 94D-54 suffered much higher stress induced by arsenate toxicity than P-deprived genotype 94D-64, in terms of lipid peroxidation, tissue H2O2 concentrations and exosmosis of K, P and As. However, P-deprived 94D-54 also had higher overproduction of enzymatic antioxidants (with higher GPX, SOD, CAT) and NPT (non-protein thiols) than P-deprived 94D-64. It appeared that, the higher sensitivity of P-deprived 94D-54 to arsenate toxicity might cause the overproduction of NPT, thus leading to the depletion of GSH and to the accumulation of H2O2. The differential sensitivity of the two genotypes has major implications for breeding rice for As affected paddy soil.
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The response of arsenate and non-tolerant Holcus lanatus L. phenotypes, where tolerance is achieved through suppression of high affinity phosphate/arsenate root uptake, was investigated under different growth regimes to investigate why there is a polymorphism in tolerance found in populations growing on uncontaminated soil. Tolerant plants screened from an arsenic uncontaminated population differed, when grown on the soil from the populations origin, from non-tolerants, in their biomass allocation under phosphate fertilization: non-tolerants put more resources into tiller production and down regulated investment in root production under phosphate fertilization while tolerants tillered less effectively and did not alter resource allocation to shoot biomass under phosphate fertilization. The two phenotypes also differed in their shoot mineral status having higher concentrations of copper, cadmium, lead and manganese, but phosphorus status differed little, suggesting tight homeostasis. The polymorphism was also widely present (40%) in other wild grass species suggesting an important ecological role for this gene that can be screened through plant root response to arsenate.
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The addition of L-Glutamate (L-GLU) and L-Hethionine ~ulfoximine (L-HSO) to mechanically isolated. photosynthetically competent, Asparagus sprengeri mesophyll cells ~u~pended in 1mM CaS04 cau~ed an immediate transient alkalinization of the cell su~pension medium in both the light and dark. The alkalinization response was specific and stereospecific as none of the L-isomers of the other 19 protein amino acids tested or D-GLU gave this response. Uptake of 14C-L-GLU was stimulated by the light. The addition of non-radioactive L-GLU. or L-GLU analogs together with 14C-L-GLU showed that only L-GLU and L-HSO stimulated alkalinization whilst inhibiting the uptake of 14C-L-GLU. Both the L-GLU dependent alkalinization and the upt~ke of 14C-L-GLU were stimulated when the external pH was decreased from 6.5 to 5.5. Increasing external K+ concentrations inhibited the uptake of 14C-L-GLU. Fusicoccin (FC) stimulated uptake. The L-GLU dependent alkalinization re~ponse exhibited monophasic saturation kinetics while the uptake of 14C-L-GLU exhibited biphasic saturation kinetics. In addition to a saturable component. the uptake kinetics also showed a linear component of uptake. Addition of L-GLU and L-MSO caused internal acidification of the cell as measured by a change in the distribution of 14C-DMO. There was no change in K+ efflux when L-GLU was added. A H+ to L-GLUinflux stoichiometry of 3:1 wa~ mea~ured at an external I.-GLU concentration of O.5mM and increased with increasing external 13 L-QLU concentration. Metabolism of L-GLU was detected manometrlcally by observing an increase in COa evolution upon the addition of L-QLU and by detection of i*C02 evolution upon the addition of »*C-L-GLU. »*C02 evolution was higher in the dark than in the light. The data are consistent with the operation of a H+/L-QLO cotransport system. The data also show that attempts to quantify the stoichlometry of the process were complicated by the metabolism of L-GLU.
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Please consult the paper edition of this thesis to read. It is available on the 5th Floor of the Library at Call Number: Z 9999.5 B63 P54 2007
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La pratique d’activité physique fait partie intégrante des recommandations médicales pour prévenir et traiter les maladies coronariennes. Suivant un programme d’entraînement structuré, serait-il possible d’améliorer la réponse à l’exercice tout en offrant une protection cardiaque au patient? C’est ce que semblent démontrer certaines études sur le préconditionnement ischémique (PCI) induit par un test d’effort maximal. Les mêmes mécanismes physiologiques induits par le PCI sont également observés lorsqu’un brassard est utilisé pour créer des cycles d’ischémie/reperfusion sur un muscle squelettique. Cette méthode est connue sous l’appellation : préconditionnement ischémique à distance (PCID). À l’autre extrémité du spectre de l’activité physique, des sportifs ont utilisé le PCDI durant leur échauffement afin d’améliorer leurs performances. C’est dans l’objectif d’étudier ces prémisses que se sont construits les projets de recherches suivants. La première étude porte sur les effets du PCID sur des efforts supra maximaux de courte durée. Les sujets (N=16) ont exécuté un test alactique (6 * 6 sec. supra maximales) suivi d’un test lactique (30 secondes supra maximales) sur ergocycle. Les sujets avaient été aléatoirement assignés à une intervention PCID ou à une intervention contrôle (CON) avant d’entreprendre les efforts. La procédure PCID consiste à effectuer quatre cycles d’ischémie de cinq minutes à l’aide d’un brassard insufflé à 50 mm Hg de plus que la pression artérielle systolique sur le bras. Les résultats de ce projet démontrent que l’intervention PCID n’a pas d’effets significatifs sur l’amélioration de performance provenant classiquement du « système anaérobie », malgré une légère hausse de la puissance maximal en faveur du PCID sur le test de Wingate de trente secondes (795 W vs 777 W) et sur le test de force-vitesse de six secondes (856 W vs 847 W). Le deuxième essai clinique avait pour objectif d’étudier les effets du PCID, selon la méthode élaborée dans le premier projet, lors d’un effort modéré de huit minutes (75 % du seuil ventilatoire) et un effort intense de huit minutes (115 % du seuil ventilatoire) sur les cinétiques de consommation d’oxygène. Nos résultats démontrent une accélération significative des cinétiques de consommation d’oxygène lors de l’intervention PCID par rapport au CON aux deux intensités d’effort (valeur de τ1 à effort modéré : 27,2 ± 4,6 secondes par rapport à 33,7 ± 6,2, p < 0,01 et intense : 29,9 ± 4,9 secondes par rapport à 33,5 ± 4,1, p < 0,001) chez les sportifs amateurs (N=15). Cela se traduit par une réduction du déficit d’oxygène en début d’effort et une atteinte plus rapide de l’état stable. Le troisième projet consistait à effectuer une revue systématique et une méta-analyse sur la thématique du préconditionnement ischémique (PCI) induit par un test d’effort chez les patients coronariens utilisant les variables provenant de l’électrocardiogramme et des paramètres d’un test d’effort. Notre recherche bibliographique a identifié 309 articles, dont 34 qui ont été inclus dans la méta-analyse, qui représente un lot de 1 053 patients. Nos analyses statistiques démontrent que dans un effort subséquent, les patients augmentent leur temps avant d’atteindre 1 mm de sous-décalage du segment ST de 91 secondes (p < 0,001); le sous-décalage maximal diminue de 0,38 mm (p < 0,01); le double produit à 1 mm de sous-décalage du segment ST augmente de 1,80 x 103 mm Hg (p < 0,001) et le temps total d’effort augmente de 50 secondes (p < 0,001). Nos projets de recherches ont favorisé l’avancement des connaissances en sciences de l’activité physique quant à l’utilisation d’un brassard comme stimulus au PCID avant un effort physique. Nous avons évalué l’effet du PCID sur différentes voies métaboliques à l’effort pour conclure que la méthode pourrait accélérer les cinétiques de consommation d’oxygène et ainsi réduire la plage du déficit d’oxygène. Nos découvertes apportent donc un éclaircissement quant à l’amélioration des performances de type contre-la-montre étudié par d’autres auteurs. De plus, nous avons établi des paramètres cliniques permettant d’évaluer le PCI induit par un test d’effort chez les patients coronariens.
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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 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.
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Phosphorus (P) deficiency is a major constraint to pearl millet (Pennisetum glaucum L.) growth on acid sandy soils of the West African Sahel. To develop cost-effective fertilization strategies for cash poor farmers, experiments with pearl millet were conducted in southwestern Niger. Treatments comprised single superphosphate hill-placed at rates of 1, 3, 5 or 7 kg P ha^−1 factorially combined with broadcast P at a rate of 13 kg ha^−1. Nitrogen was applied as calcium ammonium nitrate at rates of 30 and 45 kg ha^−1. At low soil moisture, placement of single superphosphate in immediate proximity to the seed reduced seedling emergence. Despite these negative effects on germination, P placement resulted in much faster growth of millet seedlings than did broadcast P. With P application, potassium nutrition of millet was improved and seedling nitrogen uptake increased two- to three-fold, indicating that nitrogen was not limiting early millet growth. Averaged over the 1995 and 1996 cropping seasons, placed applications of 3, 5 and 7 kg P ha^−1 led to 72%, 81% and 88% respectively, of the grain yield produced by broadcasting 13 kg P ha^−1. Nitrogen application did not show major effects on grain yield unless P requirements were met. A simple economic analysis revealed that the profitability of P application, defined as additional income per unit of fertilizer, was highest for P placement at 3 and 5 kg ha^−1.