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  • About
  • The Global ETD Search service is a free service for researchers to find electronic theses and dissertations. This service is provided by the Networked Digital Library of Theses and Dissertations.
    Our metadata is collected from universities around the world. If you manage a university/consortium/country archive and want to be added, details can be found on the NDLTD website.
121

Water stress effects on growth, yield and quality of wheat (Triticum aestivum L.)

Mbave, Zwidofhelangani Aubrey 25 April 2013 (has links)
Understanding the effects of water stress on wheat growth, yield and quality is essential for good irrigation management. In South Africa most of the wheat production areas are vulnerable to drought stress during crop development. That causes substantial reduction in grain yield, depending on the developmental stage at which water stress occurred. Supplemental irrigation is the main strategy for adaptation and stabilisation of yield under water stress. However, agriculture is the leading single water-use sector locally, consuming about 60% of total available water. Therefore, the need to improve water use efficiency (WUE) in crop production is clear, since South Africa is classified as a water-scarce country. Experiments were conducted under a rain shelter at Hatfield Experimental Farm, University of Pretoria, in the 2010 and 2011 seasons. The main objective of the study was to evaluate the effects of water stress at different stages on growth, yield, and quality of three wheat cultivars, namely Duzi, Steenbras and SST 843. Water stress was imposed by withholding water at either of three growing stages. The first treatment was stressed during tillering stages to flag leaf (stem elongation (SNN)), followed by water stress from flag leaf to the end of flowering (flowering stage (NSN)), and lastly water stress from grain filling to physiological maturing (grain-filling stage (NNS)), whereas optimal supply of water was maintained throughout the season by weekly irrigating to field capacity for the control treatment (NNN). Irrigation treatments and cultivars influenced growth, yield and quality, depending on the developmental stage at which irrigation was withheld. The control treatment (NNN) and the treatment stressed in the flowering stage (NSN) had highest and lowest grain yield respectively in both seasons. Water stressed treatment NSN reduced grain yield by 33% and 35% in the 2010 and 2011 seasons respectively, when compared with the control treatment (NNN). Reduction of grain yield due to stress in the flowering stage (NSN) was ascribed to reduction in the number of seeds per ear, number of ears per unit area, ear length, and flag-leaf photosynthesis rate (Pn). In the flowering stage (NSN) water stress reduced Pn by 59% which was due to increased leaf temperature because of lower transpiration (E) and stomatal conductance (gs). The water stress treatment NSN reduced transpiration by 72% and stomatal conductance by 84% in the flowering stage. Plant height was reduced by 23% because of water stress imposed in the flowering stage (NSN), which consequently decreased biomass yield by 29% in the 2011 season. Growth and yield parameters showed dramatic recovery when stress was terminated during the flag-leaf stage (SNN). The cultivar Steenbras had lower yield reduction under stress, whereas Duzi and SST 843 had higher yield potential under the well-watered conditions (NNN). In the 2011 season SST 843 had higher WUE of 14.2 kg ha-1 mm, which corresponded to higher grain yield of 7210 kg ha-1 and higher ET of 509 mm. Water-stress treatment SNN gave the highest WUE of 14.9 kg ha-1 mm, which corresponded to a total water use (ET) of 451 mm and grain yield of 6738 kg ha-1. Water stress treatments SNN and NNS reduced ET by 27% and 17%, respectively, which translated to 173 mm and 105 mm water saved by each treatment correspondingly. Grain protein content (GPC) was reduced most by the treatment exposed to stress in the stem elongation stage (SNN). However, the GPC was acceptable (>12%) in all treatments in both seasons. Hectolitre mass was reduced most by water stress imposed during grain filling (NNS). Water stress treatment NNS lowered the hectolitre mass by 3% and 4% in the 2010 and 2011 seasons respectively. Generally all quality parameters in the present study were acceptable for all irrigation treatment and cultivars. The hypothesis that water stress in the stem elongation and grain-filling stages will have little effect on yield and improve WUE was accepted. Therefore it can be recommended that supplemental irrigation should be applied from flag leaf to end of flowering (NSN) stages of wheat in order to minimise grain yield losses in the absence of rainfall. Further research should focus on extrapolation of these results to other production regions using crop models. / Dissertation (MInstAgrar)--University of Pretoria, 2013. / Plant Production and Soil Science / unrestricted
122

Impact du déficit hydrique sur la dégradabilité, la biochimie pariétale et la répartition des tissus lignifiés chez l’entrenoeud de maïs et déterminisme génétique de ces caractères / Impact of water deficit on degradability, cell wall biochemistry and lignified tissue distribution in the maize internode, and genetic determinism of these traits

El Hage, Fadi 20 December 2018 (has links)
Ce projet de thèse s’inscrit dans un contexte de changement climatique et de remplacement des énergies fossiles, où la réduction des apports en eau et l’optimisation de la valorisation de la biomasse sont deux enjeux majeurs des systèmes de productions durables. La dégradabilité de la biomasse est principalement limitée par la dégradabilité des parois et afin de l’améliorer, il est nécessaire de comprendre quels facteurs sont impliqués dans cette limitation de dégradabilité. Plusieurs études ont montré que la dégradabilité pariétale est impactée par la composition et la structure de la paroi mais aussi par la distribution des tissus lignifiés au sein des organes. Pour faire la part entre l’impact de la biochimie et celui de l’histologie sur la dégradabilité dans différentes conditions d’irrigation, des outils haut-débit de quantifications biochimiques et histologiques ont été développés et dédiés à l’étude d’entrenoeuds portant l’épi principal. Les études ont porté sur un panel de diversité génétique de maïs et d’une population de lignées recombinantes, cultivés durant plusieurs années dans des conditions d’irrigation contrastées dans le sud de la France. Nos résultats mettent en évidence que le déficit hydrique induit une augmentation de la dégradabilité pariétale, accompagnée par une diminution de la teneur en lignines pariétales et par une induction préférentielle d’une lignification corticale, plus p-coumaroylée. De façon originale, nous avons aussi cartographié 90 QTLs de caractères histologiques dans les différentes conditions d’irrigation sur le génome du maïs. Plus particulièrement, une large région entre le bin 1.07 et le bin 1.11 est impliquée dans les variations observées du nombre de faisceaux vasculaires et de la surface de section des entrenoeuds. De façon globale, de nombreux QTLs de composition pariétale de l’entrenoeud co-localisent avec ceux obtenus au niveau de la plante entière sans épis chez la même population. Enfin nous avons pu démontrer que le choix de l’entrenoeud portant l’épi principal est judicieux pour représenter à la fois les caractéristiques histologiques des tiges entières et biochimiques de la biomasse lignocellulosique des plantes entières. Ainsi, les caractéristiques histologiques et biochimiques des entrenoeuds de maïs sont proposées comme des cibles de choix pour sélectionner des lignées de maïs résilientes au déficit hydrique. / This PhD project encompass in a context of global warming and replacement of fossil resources, where both water supply decrease and biomass valorisation optimisation are two major issues for providing sustainable systems of production. Biomass degradability is mainly limited by cell wall degradability and in order to improve it, it is necessary to understand what are the factors involved in the limitation of the degradability. Several studies have shown that cell wall degradability is impacted by the structure and the composition of the cell wall but also by the distribution of the lignified tissues within the organs. To decipher the impact of the biochemistry and from the one of the histology on degradability under different watering conditions, high-throughput quantifications tools for histology and biochemistry have been developed and dedicated to the study of the internode carrying the main ear. The studies were conducted on a maize genetic diversity panel and a recombinant inbred lines population, cultivated during several years under contrasted irrigation conditions in South of France. Our results highlight that water deficit induce an increase of the cell wall degradability, associated to a decrease of the cell wall lignin content and a preferential induction of a cortical lignification, more p-coumaroylated. In original ways, we also detected 90 QTLs for histological traits under the different irrigations scenarios on the maize genome. More particularly, a large region between bin 1.07 and bin 1.11 is involved in the observed variations of the bundle number and the stem section area of the internodes. More generally, numerous QTLs of cell wall composition of the internode co-localised with the ones detected for the whole plant without ear level in the same population. Finally we were able to demonstrate that the choice of the internode carrying the main ear is judicious to represent both histological characteristics from the whole stem and biochemical characteristics from the lignocellulosic biomass of the whole plant. Thus, histological and biochemical traits in maize internode are proposed as particular targets to select lines resilient to water deficit.
123

Determinants of sunflower seed quality for processing

Nel, Andries Abraham 01 September 2001 (has links)
The low and varying protein content and high crude fibre content of sunflower oil cake produced from sunflower seed create problems for the South African oil expelling industry. This prompted research into factors that may affect the seed quality for processing purposes. The seed quality characteristics are the seed oil and protein contents and the hullability. Analysis of the kernel-rich fraction produced after dehulling gives an indication of the potential oil yield, oil cake yield and oil cake protein and crude fibre contents and thus the processed value. Seed hullability and potential losses of oil and protein were affected by seed moisture content and seed size. Drying seed resulted in increased hullability, and sifting it into size classes proved to be a mechanism for differentiating in terms of oil cake quality. The effects of cultivar, environment and selected environmental variables on seed yield and processing quality were investigated by means of field trials. Seed yield and quality were more affected by environment than by cultivar. Seed size and hullability, and as a result also the protein content of the potential oil cake, were affected by plant population, with lower populations associated with better quality. Increased nitrogen application improved seed yield and seed protein content but lowered seed oil content, with no effect on hullability. Boron fertilisation improved seed yield in one trial but suppressed yield in a second trial. Hullability declined in one trial due to boron fertilisation. A mild water stress during the grain-filling stage reduced seed yield by 23% and hullability by 14%. Optimising the seed oil:seed protein ration through breeding may be the most advisable option for improving seed quality for processing. Due to the need for a seed grading system based on seed quality, regression analyses between easily measurable seed characteristics and seed quality parameters were done. The relatively low mean deviation between measured and predicted values indicate that seed oil content, protein content and hullability can be estimated with reasonable accuracy. These relationships must still be validated. / Dissertation (PhD (Plant Production: Agronomy))--University of Pretoria, 2002. / Plant Production and Soil Science / unrestricted
124

Stanovení těkavých izoprenoidů jako markerů vlivu vodního stresu na rezistenci smrku vůči kůrovcům / Determination of volatile isoprenoids as water stress markers of spruce resistance against bark beetle

Slušná, Michaela January 2012 (has links)
Isoprenoids are important components of conifer resin and represent an important part of constituted defence system against herbivores and pathogens. Drought is one of the most important factors that influences the tree physiology and resitance. Due to decreased turgor of resin canal cells, the water insufficiency affects the pressure of the resin and thereby the ability of trees to physically prevent pathogen or herbivore invasion by effective outpouring of the resin. In addition, drought can also change the resin composition and thus can influence the quality of volatiles emitted by the tree. The Norway spruce, Picea abies, is the predominant species of production forests in moderate climate zone. Bark beetles, Ips typographus, represent the most important pest species of spruce. In general, pioneer bark beetles use host volatiles to orient themselves toward the tree suitable for colonization and in many species host volatiles synergize bark beetle aggregation pheromones. Thus the host volatile composition could affect significantly host colonization. This diploma thesis studied the influence of drought on the production and composition of isoprenoid volatile organic compounds in 80 - 100 years old spruce trees. Using I. typographus antennae as biological detectors, we also studied which resin...
125

Community Interactions and Water as Drivers of Soil Microbial Communities

Kakumanu, Madhavi Latha 06 August 2011 (has links)
Understanding the response of soil microbial communities to various environmental stresses is of current interest, because of their pivotal role in nutrient cycling, soil organic matter mineralization and influence on plant growth. Determining the affect of several biotic and abiotic factors on soil microbial communities is the overall objective of the study. The specific goals are to determine 1) the response of microbial communities to water deficit in soil and 2) how the presence of a rich biotic community determines the direction of microbial community development in cultures. Both goals are novel and unique contributions to understanding microbial ecology in soil. Dynamics in water potentials due to drying and rewetting of soil impose significant physiological challenges to soil microorganisms. To cope with these fluctuations, many microorganisms alter the chemistry and concentration of their cytoplasmic contents. The aim of this research is to understand how the microbial biomass and their cytoplasm change in response to water potential deficits under in situ soil conditions. To address this objective we characterized intracellular and extracellular metabolites in moist, dry and salt stressed soils. Our results provided the first direct evidence that microbial communities in soil in situ utilize sugars and sugar alcohols to cope with low water potential. While the cultivation and isolation of microorganisms is essential to completely explore their physiology and ecology, 99% of soil microbes resist growing in cultures. Presence of very unnatural conditions in the culture plates was considered as main reason for low cultivability. Thus, a culture-based study was conducted whereby microorganisms were grown in association with their native habitat with an objective of mimicking native conditions to promote the growth of previously uncultivated microorganisms. Moreover, the importance of biotic communities (microbe-microbe) and abiotic soil effects were assessed on bacterial growth. Our results strongly indicate that the presence of living microbial community in the vicinity of the target culture resulted in the cultivation of novel members of rare bacterial taxa from phyla Verrucomicrobia, Bacteroidetes, Proteobacteria, and Planctomycetes. These results emphasize the need to develop new culturing methods to tap the hidden microbial potential for emerging anthropogenic needs.
126

Characterization of a Novel Water Stress Related Protein from Geranium (<em>Pelargonium hortorum</em>).

Ashley, David W 01 December 2001 (has links) (PDF)
Searches for sequence similarity against a previously isolated and known ABA regulated partial geranium cDNA on GenBank and TIGR databases gave results indicating sequence similarities to both firefly luciferase and 4-coumarate:CoA Ligase (4-CL). To determine whether or not this geranium cDNA, and a root mRNA from Arabidopsis that hybridized with the geranium cDNA, is 4-CL, specific Arabidopsis 4-CL primers were used for expression analysis by RT-PCR of Arabidopsis 4-CL mRNA from Arabidopsis root tissue. The expression pattern of the Arabidopsis 4-CL mRNA was identical with the expression pattern of the mRNA that hybridized with the geranium cDNA. This matching expression pattern gives very strong evidence that our geranium cDNA is 4-CL like and that Arabidopsis 4-CL is downregulated during water stress by ABA. This is significant in that it has not previously been reported. Northern analysis and PCR product sequence analysis further confirmed this expression data.
127

Landscape performance of Callistemon citrinus under environmental stress conditions

Mohsin, Riyadh Mannaa 09 August 2019 (has links)
Woody ornamental plants are considered the key for well- designed landscapes; Callistemon citrinus is one of those plants. In arid and semiarid areas, Callistemon has been used in gardens and landscapes for its unique characteristics. This study was conducted to evaluate the performance of Callistemon in landscapes under different environmental stress conditions including water, and heat. Further, the potential of using 3D software SketchUp in landscape design was also investigated. Callistemon plants were grown under 100, 50 or 30% of field capacity with or without shredded hardwood mulch. When field capacity decreased, root: shoot ratio and water use efficiency were improved, but canopy temperature increased. The decreased water caused decreases in leaf greenness, chlorophyll a, shoot volume, root volume, leaf water potential, and transpiration rate. Mulch increased root volume and leaf water potential. Callistemon growth under shade structure angle and orientation was examined. Shade structures were installed at 90° and 70° to the ground and three orientations, south, east, and west. Transpiration rate was reduced under the shade. The plant’s response to heat was also examined. Plant were exposed to 45/35, 35/25, or 25/15 °C, d/n. The plant’s physiological response was tested after two weeks. Highest temperatures decreased Chl b content and SOD activity compared to control. In contrast, carotenoid content and H2O2 level increased under the highest temperature treatment. Catalase activity was increased at moderate but decreased under the highest temperatures. Designing a 3D model using SketchUp software was examined. The program function was evaluated, and the designed experiment was tested. Planning a site with shadows predicted was achieved. The work was done with greater precision and less effort. The program was effective in reducing time and cost. SketchUp can be successfully used in landscape work.
128

The combined effects of fertilization and relative water limitation on tissue water relations, hydraulic parameters and shallow root distribution in loblolly pine (Pinus taeda L.)

Russell, Edward Morgan 27 August 2019 (has links)
One goal of this research was to characterize shoot tissue-level responses in loblolly pine to soil moisture limitation in combination with fertilization as well as to more severe soil moisture limitation. We found that neither fertilization alone, nor fertilization in combination with soil moisture limitation resulted in changes to shoot tissue water relations parameters classically characterized in drought response studies. More severe water limitation was necessary to elicit responses, and those responses had not been fully described previously. The more severe water limitation resulted in increased capacitance beyond turgor loss, increased relative water content at turgor loss, a more negative turgor loss point, an increased bulk modulus of elasticity, more negative osmotic potential at 100% relative water content, and an increased apoplastic water fraction. As there were indications of reduced water use and moisture stress in the absence of shoot level responses under less severe drought, such parameters are insufficient alone to characterize moisture stress in fertilized and in less severely water limited loblolly trees. Additionally, we sought a morphological or physiological explanation for the reduced transpiration and increased water use efficiency reported for fertilized trees in the Virginia Piedmont. Our characterizations of the responses of root distribution and hydraulics to limited soil moisture here complement existing research, which demonstrated changes to root distribution and hydraulics in response to fertilization. The responses we discovered in fertilized trees that accompanied reduced transpiration and increased water use efficiency that differed from responses to reduced soil moisture alone were primarily large decreases to shallow root presence. We found this to be readily quantified using measures of root length density. Decreases to whole-tree hydraulic conductivity were also shown to occur with fertilization and were shown not to occur in shoot tissue, suggesting limitation via rhizosphere or root xylem conductance. Our results support the supposition that fertilization narrows hydraulic safety margins and potentially predisposes loblolly trees to moisture stress, particularly prolonged, severe water limitation following fertilization. Finally, we tested the validity of throughfall exclusion for simulating reduced rainfall using a greenhouse 'split-pot' study, which applied spatially fixed heterogeneous soil moisture to young, well-watered loblolly pines. The 'split-pot' experiments demonstrated that spatially fixed soil moisture heterogeneity does not confound drought effects; needle area specific transpiration was not decreased, nor was water use efficiency increased. This supports the validity of inferences taken from drought simulation experiments with loblolly pine where throughfall exclusion troughs reduce soil moisture content in a consistent, spatially heterogeneous manner. / Doctor of Philosophy / We investigated various effects of soil moisture limitation alone, and in combination with common fertilization practices in loblolly pine production. Responses at the shoot and needle level to different levels of soil moisture limitation produced new findings concerning how tissues respond to more severe water limitation. A 30% decrease in throughfall precipitation alone, or in combination with fertilization did not elicit drought related shoot tissue responses despite the presence of other indications of moisture stress and reduced water use. We also sought to explain why fertilized trees experiencing water limitation had environmental sensitivities that were different from unfertilized tree receiving ambient rainfall amounts or from trees only experiencing water limitation without fertilization. We found that changes to shallow root presence, especially root length density, accompanied the different patterns of environmental sensitivity and water use. Also, the water conducting ability of roots changed unevenly in soil with uneven moisture levels. The ability of roots to resist loss of conductivity to water did not change unevenly in the same way. We did another set of experiments to determine if using impervious troughs to catch rain is a valid approach to reducing soil moisture for the purpose of testing how loblolly responds to water limitation. These throughfall exclusion troughs create uneven soil moisture reduction, which can have effects on plant water use that are separate from water limitation alone. We found that in well-watered young trees, uneven soil moisture alone did not produce responses that could be confused with the effects of water limitation. This finding indirectly validates the use of throughfall exclusion troughs to simulate reduced rainfall.
129

Exploring the Soil-Plant-Atmosphere Continuum: Advancements, Integrated Modeling and Ecohydrological Insights

D'Amato, Concetta 31 May 2024 (has links)
In recent years, the Soil-Plant-Atmosphere (SPA) continuum has faced unprecedented challenges due to anthropogenic modifications and climate change. Understanding the complex dynamics of this system in response to such changes is crucial for addressing contemporary environmental concerns. Albert Einstein's famous quote, "The measure of intelligence is the ability to change", resonates deeply throughout this doctoral thesis. This thesis aims to address the complex issue of SPA interactions by developing a comprehensive set of models capable of representing the intricate dynamics of this system. At the core of this research lies the integration of sophisticated descriptions of hydrological and plant biochemical processes into a novel ecohydrological model, GEOSPACE-1D (Soil Plant Atmosphere Continuum Estimator model in GEOframe). Through a combination of theoretical exploration, engineering methodologies, and empirical experiments, this thesis aims to advance our understanding of SPA interactions. The development of adaptable models, represents a significant contribution to the field. The thesis emphasizes the practical implications of employing models to analyze experimental data, thereby enhancing our comprehension of various phenomena. In conclusion, this thesis provides valuable insights into SPA interactions and lays the groundwork for future research and applications. By embracing the challenge of understanding and modeling the SPA continuum, this work contributes to the ongoing efforts to address environmental challenges and promote sustainable practices.
130

Impacts des défoliations de la processionnaire du pin (Thaumetopoea pityocampa) sur la croissance du pin maritime (Pinus pinaster)

Jacquet, Jean-Sébastien 17 December 2012 (has links)
Parmi les perturbations attendues des écosystèmes forestiers dans le cadre du changement climatique, les dégâts causés par les insectes ravageurs devraient augmenter en raison notamment de la hausse des températures. Ces dommages devraient réduire la capacité des forêts à stocker du carbone, et donc à atténuer les causes du changement climatique. Plus spécifiquement, il est prédit que l’augmentation des pullulations d’insectes défoliateurs conduise à une diminution importante de la productivité des forêts mais leur impact réel est difficile à quantifier et à comprendre. L’objectif général de ce travail a donc été d’améliorer nos connaissances sur l’impact d’une défoliation sur la croissance d’arbres d’âges contrastés et soumis à différentes contraintes environnementales. Dans un premier temps, ce travail a permis de quantifier la relation entre l’intensité de défoliation de la processionnaire du pin et la perte de production du pin maritime. Une méta-analyse nous a permis de synthétiser les résultats de l’ensemble des 45 études publiées à ce jour sur le sujet. Par la suite, profitant d’une pullulation exceptionnelle de processionnaire du pin pendant l’hiver 2009-2010, nous avons expérimentalement testé l’effet de défoliations variant de 25 à 100% en interaction avec l’âge des peuplements de pin maritime. Nous concluons que les pertes de production sont proportionnelles à l’intensité de défoliation et augmentent avec l’âge de l’arbre.L’évolution du climat étant susceptible de favoriser la combinaison de différents stress, nous nous sommes également intéressés aux effets cumulés de la défoliation et du stress hydrique sur la croissance de l’arbre. Notre étude révèle que la défoliation et le stress hydrique ont des effets additifs sur la croissance du pin maritime. Nos résultats confirment la complexité de la réponse éco-physiologique de l’arbre à la défoliation. L’étude des réserves carbonées et azotées nous a permis d’explorer les possibles mécanismes sous-jacents à l’effet d’une défoliation. Plus qu’une simple diminution de la fixation carbonée par altération de l’appareil photosynthétique, nous proposons que la défoliation affecte la croissance via des processus de limitation en ressources carbonées et azotées. Les réserves carbonées n’étant affectées qu’en début de saison, nous émettons l’hypothèse d’une limitation azotée de l’arbre après défoliation. Ainsi, la défoliation et le stress hydrique pourraient diminuer conjointement la disponibilité en eau et en ressources minérales, ce qui expliquerait leurs effets additifs sur la croissance de l’arbre. / In the context of climate change, forest pest outbreaks, among other biotic disturbances, are expected to be more frequent in response to increasing temperatures. The resulting damage is likely to adversely affect forests net primary production and their contribution to climate mitigation via carbon sequestration. More specifically, insect defoliators are to predicted to be more prevalent in the future but their real impact on forest productivity is difficult to evaluate and interpret. Our main objective was then to improve our understanding of insect defoliation impact on tree growth at different tree ages and under various climatic conditions.First, we assessed the relationship between the intensity of pine processionary moth defoliation and maritime pine growth loss. We carried out a meta-analysis to summarize the outcomes of the 45 published studies that addressed this issue. Then, we took advantage of a severe pine processionary moth outbreak to set up a large field experiment where we controlled for both the age (from 3 to 40 years old) and the defoliation rate (from 25 to 100%) of Pinus pinaster trees. We showed that radial growth losses were proportional to defoliation intensity and more important in older trees.As the combination of several disturbances is likely to be more frequent under climate change, we developed a manipulative experiment to quantify the cumulative impact of water stress and defoliation on maritime pine tree growth. We found additive detrimental effects of water stress and defoliation on maritime pine tree growth.Our results confirm that tree response to defoliation, including various eco-physiological processes, is complex. To decipher the underlying mechanisms we analyzed the dynamics of nutrient and carbohydrates pools in defoliated trees during the growing season. Our findings suggest that defoliation affect tree growth through resource limitation rather than via a reduction of carbon fixation due to altered photosynthesis. Since carbohydrates pools were only affected early in the season, our results support the hypothesis of a nitrogen limitation in trees following defoliation. Additive effects of defoliation and water stress may then be explained by similar adverse consequences on water flow and nitrogen uptake.

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