The nature and occurrence of mycorrhizal associations, with particular reference to the anatomy, carbohydrate physiology, plant mineral nutrition and occurrence of ectomycorrhizae (ECM), is discussed. The ecology and forest relations of <i>Picea sitchensis</i> - the Sitka spruce concludes the literature review. Identification of areas of good and poor Sitka growth on related soils and the quantification of their ECM status, investigation of the effect of phosphate addition to Sitka seedlings in pots, subsequent and changes to their ECM status, and the effects of soluble aluminium on phosphate nutrition of Sitka seedlings, the collation of results and relation back to the field situation were carried out as experimental work. Field sites were identified and described in terms of geology, soils, field ECM status, forest productivity and nutrient status. Three pot experiments were carried out. The 1st investigated the effects of phosphate application on ECM Sitka seedlings in soil from the field sites; the 2nd investigated the effects of phosphate application to ECM and non-mycorrhizal (NM) seedlings in compost; and the 3rd investigated the affects of application of Al-citrate to ECM and NM seedlings in compost containing high and low levels of phosphate. The results obtained were described and discussed in the context of a model of the factors affecting plant response to the soil environment. The field ECM development representing a considerable drain on the carbohydrate economy of the field sites was to some extent duplicated in the greenhouse. The possible decrease in importance of this drain was illustrated by phosphate application, but was increased by addition of Al-citrate. A previously unrecorded ECM-enhanced uptake of Manganese was reported. The importance of phosphate in the soils under discussion was emphasised, and possible further work suggested.

Tropical soils are a globally important store of terrestrial carbon (C) and source of atmospheric carbon dioxide (CO2), regulated by the activity of soil microorganisms, through the mineralisation of plant residues and soil organic matter (SOM). Climatic warming will influence microbial activity, and this may accelerate the rate of C release from soils as CO2, contributing to alterations in current atmospheric composition, and generating feedbacks to climate change. Yet the magnitude of C loss from tropical soils remains uncertain, partly because we do not fully understand how non-climatic factors – including the chemistry of plant inputs, the availability of soil nutrients and the composition of the decomposer community – will interact to determine the response to changes in temperature. This thesis examines how these factors together regulate the rate of C cycling in contrasting soils across a 3400 m tropical elevation gradient in the Peruvian Andes, spanning a 20 ºC range (6.5 – 26.4 ºC) in mean annual temperature. Large-scale field-based manipulation experiments, translocating leaves and soil-cores across the elevation gradient (to impose an in-situ experimental warming treatment), were combined with controlled laboratory studies to examine the microbial-scale mechanisms which underlie the processes of decomposition and soil respiration observed in-situ. Results show that, across the gradient, rates of leaf-decomposition were determined principally by temperature and foliar chemical traits, while soil fertility had no significant influence. The effect of temperature was, however, stronger across higher-elevation sites, suggesting a greater vulnerability of the C-rich soils in montane systems to increased C loss under climatic warming. In lowland forests, the presence of invertebrate macrofauna also accelerated rates of decomposition, but leaf chemistry explained the greatest proportion of the observed variance, with a strong role for leaf chemical traits also identified under controlled conditions. Despite marked differences in microbial abundance and community composition among soils, these metrics were not associated with observed rates of decomposition. These results suggest that climate-related changes to plant species distributions (with associated changes to the chemistry of leaf-inputs), and upslope extension of macrofaunal ranges, could strongly influence future rates of leaf decomposition, independently of the direct response to warming. From the soil translocation study, root-soil interactions stimulated substantial net C loss from montane soils following translocation downslope (experimental warming treatment), indicating that warming-related changes to root productivity, exudation and/or species-composition could represent an important mode of future C loss from these soils. To examine more closely how inputs of plant-derived C influence the turnover of pre-existing SOM, and whether soil nutrient availability modulates the response, soils were amended with simple and complex 13C-labelled substrates in combination with inorganic nutrient treatments. Isotopic partitioning was used to determine the degree to which C and nutrient inputs accelerated (positive priming) or retarded (negative priming) the decomposition of SOM. Amendment of upper montane forest and montane grassland soils with nitrogen (N; alone and in combination with C) substantially retarded the decomposition of SOM, suggesting that microbial demand for N strongly regulates the turnover of organic matter in these soils. In contrast, amendment of lower montane and lowland forest soils with C stimulated positive priming of SOM, which was strongest in response to the simple C substrate and was not influenced by nutrient treatments, suggesting that microorganisms in these soils are primarily constrained by availability of labile C. Functional differences among microbial groups were also evident, with gram-negative bacteria and fungi using more labile sources of C while gram-positive bacteria used more complex C. Together, results from these studies considerably advance our understanding of soil C dynamics across lowland and montane systems, painting a rich picture of interacting processes which will determine the future soil C balance in tropical ecosystems. They show that the influence of temperature on the rate of soil C cycling is strongly affected by the nature and composition of plant-derived and atmospheric inputs, the principal additional constraints varying with elevation, leading to both opposing and reinforcing effects on rates of decomposition. The greater observed temperature sensitivity of decomposition at higher elevations is coupled with high microbial demand for N which regulates the turnover of SOM, whereas at lower elevations leaf decomposition is accelerated by active macrofaunal breakdown, while microbial decomposition of SOM is constrained by the availability of labile C. Under a global change scenario of increased temperature and N deposition, results therefore suggest that: (i) modified chemistry of plant inputs will influence rates of decomposition, independently of climate; (ii) increased availability of labile C will lead to more rapid decomposition of SOM at lower elevations; (iii) greater root productivity (associated with warming and plant-community shifts) will stimulate soil C loss across montane regions; but (iv) at higher elevations, a possible countervailing effect may be imposed on rapid warming-accelerated decomposition if increased N availability reduces microbial mineralisation of SOM. The net effect on the ecosystem C budget will depend on the balance of C gain from primary productivity and C loss from soils. Overall, however, the results presented here suggest that the large soil C stores in higher-elevation montane regions are particularly vulnerable to substantial reductions under exposure to short- and medium-term climatic warming.

Reindeer appear to have strong positive effects on plant productivity and nutrient cycling in strongly nutrient-limited ecosystems. While the direct effects of grazing on vegetation composition have been intensively studied, much less is known about the indirect effect of grazing on plant-soil interactions. This thesis investigated the indirect effects of ungulate grazing on arctic plant communities via soil nutrient availability and plant nutrient uptake. At high density, the deposition of dung alone increased plant productivity both in nutrient rich and nutrient poor tundra habitats without causing major changes in soil possesses. Plant community responses to dung addition was slow, with a delay of at least some years. By contrast, a 15N-urea tracer study revealed that nutrients from reindeer urine could be rapidly incorporated into arctic plant tissues. Soil and microbial N pools only sequestered small proportions of the tracer. This thesis therefore suggests a strong effect of dung and urine on plant productivity by directly providing nutrient-rich resources, rather than by stimulating soil microbial activities, N mineralization and ultimately increasing soil nutrient availability. Further, defoliation alone did not induce compensatory growth, but resulted in plants with higher nutrient contents. This grazing-induced increase in plant quality could drive the high N cycling in arctic secondary grasslands by providing litter of a better quality to the belowground system and thus increase organic matter decomposition and enhance soil nutrient availability. Finally, a 15N natural abundance study revealed that intense reindeer grazing influences how plants are taking up their nutrients and thus decreased plant N partitioning among coexisting plant species. Taken together these results demonstrate the central role of dung and urine and grazing-induced changes in plant quality for plant productivity. Soil nutrient concentrations alone do not reveal nutrient availability for plants since reindeer have a strong influence on how plants are taking up their nutrients. This thesis highlights that both direct and indirect effects of reindeer grazing are strong determinants of tundra ecosystem functioning. Therefore, their complex influence on the aboveground and belowground linkages should be integrated in future work on tundra ecosystem N dynamic.

Neste trabalho é apresentada uma formulação híbrida do Método dos Elementos de Contorno/Método dos Elementos finitos à análise da interação placa-estaca-solo. Nesta formulação a placa é modelada pelo método dos elementos finitos, utilizando os elementos DKT (Discrete Kirchhoff Theory) e HSM (Hybrid Stress Model); e o solo é modelado pelo método dos elementos de contorno como um meio elástico semi-infinito. A estaca é representada por apenas um elemento, com 3 pontos nodais definidos ao longo de seu fuste e a tensão de cisalhamento ao longo da mesma é aproximada por um polinômio do segundo grau. A tensão normal à seção da extremidade inferior da estaca é suposta constante e um ponto nodal é definido no centro desta seção. A interface placa-solo é dividida em elementos de contorno triangulares coincidentes com a divisão dos elementos finitos da placa; e admite-se que a tensão de contato varie linearmente no domínio de cada elemento. As tensões de contato são eliminadas nos dois sistemas de equações, provenientes do MEF e do MEC, para obter o sistema final de equações governantes do problema. Após a resolução deste sistema são obtidos os deslocamentos nos nós e a partir deles são calculadas as tensões de contato placa-solo e a carga absorvida por cada estaca. Além disso, esta formulação também permite a análise de blocos de estacas com ou sem contato entre o bloco e o solo. Vários exemplos envolvendo a interação placa-solo, estaca-solo e placa-estaca-solo foram analisados e os resultados obtidos estão de acordo com os fornecidos por outras formulações. / This work presents a hybrid Finite Element/Boundary Element formulation for the analysis of plate-pile-soil interactions problems. In this formulation the plate is modeled by finite elements using DKT (Discrete Kirchhoff Theory) and HSM (Hybrid Stress Model) elements and the soil is modeled as an elastic half space by the boundary element method. The pile is represented by only one element, with 3 nodal points, and shear force along the pile is approximated by a second degree polynomial. The pile-tip stress is assumed to be constant over the cross-section and a further nodal point is located there. The interface plate-soil is divided in a triangular boundary elements mesh coincident with that used in the finite element of the plate and the subgrade traction is assumed to vary linearly across each element. The subgrade tractions are eliminated from the two system of equation obtained with FEM and BEM resulting in the governing system of equation for plate-pile-soil interactions problems. By solving this set of equations the nodal displacements, the load on the piles and the subgrade tractions are calculated. Besides, this formulation allows also analysis several piles groups with and without ground-contacting rigid and flexible caps. Numerical results are presented for plate-soil, pile-soil and plate-pile-soil interactions. In all the results agree closely with those from much more elaborate analysis.

Neste trabalho é apresentada uma formulação híbrida do Método dos Elementos de Contorno/Método dos Elementos finitos à análise da interação placa-estaca-solo. Nesta formulação a placa é modelada pelo método dos elementos finitos, utilizando os elementos DKT (Discrete Kirchhoff Theory) e HSM (Hybrid Stress Model); e o solo é modelado pelo método dos elementos de contorno como um meio elástico semi-infinito. A estaca é representada por apenas um elemento, com 3 pontos nodais definidos ao longo de seu fuste e a tensão de cisalhamento ao longo da mesma é aproximada por um polinômio do segundo grau. A tensão normal à seção da extremidade inferior da estaca é suposta constante e um ponto nodal é definido no centro desta seção. A interface placa-solo é dividida em elementos de contorno triangulares coincidentes com a divisão dos elementos finitos da placa; e admite-se que a tensão de contato varie linearmente no domínio de cada elemento. As tensões de contato são eliminadas nos dois sistemas de equações, provenientes do MEF e do MEC, para obter o sistema final de equações governantes do problema. Após a resolução deste sistema são obtidos os deslocamentos nos nós e a partir deles são calculadas as tensões de contato placa-solo e a carga absorvida por cada estaca. Além disso, esta formulação também permite a análise de blocos de estacas com ou sem contato entre o bloco e o solo. Vários exemplos envolvendo a interação placa-solo, estaca-solo e placa-estaca-solo foram analisados e os resultados obtidos estão de acordo com os fornecidos por outras formulações. / This work presents a hybrid Finite Element/Boundary Element formulation for the analysis of plate-pile-soil interactions problems. In this formulation the plate is modeled by finite elements using DKT (Discrete Kirchhoff Theory) and HSM (Hybrid Stress Model) elements and the soil is modeled as an elastic half space by the boundary element method. The pile is represented by only one element, with 3 nodal points, and shear force along the pile is approximated by a second degree polynomial. The pile-tip stress is assumed to be constant over the cross-section and a further nodal point is located there. The interface plate-soil is divided in a triangular boundary elements mesh coincident with that used in the finite element of the plate and the subgrade traction is assumed to vary linearly across each element. The subgrade tractions are eliminated from the two system of equation obtained with FEM and BEM resulting in the governing system of equation for plate-pile-soil interactions problems. By solving this set of equations the nodal displacements, the load on the piles and the subgrade tractions are calculated. Besides, this formulation allows also analysis several piles groups with and without ground-contacting rigid and flexible caps. Numerical results are presented for plate-soil, pile-soil and plate-pile-soil interactions. In all the results agree closely with those from much more elaborate analysis.