883 resultados para Size structure


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In living bodies, the correct perceptual representation of size constancy requires that an object's size appear the same when it changes its location with respect to the observer. At the same time, it is necessary that objects at different locations appear to be the same size if they are. In order to do that, the perceptual system must recover from the stimuli impinging on the individual, from the light falling on the retina, a representation of the relative sizes of objects in the environment. Moreover, at the same time, image perception is related to another type of phenomena. It corresponds to the well known perceptual illusions. To analyze this facts, we propose a system based on a particular arrays of receptive points composed by optical fibers and dummy fibers. The structure is based on the first layers of the mammalians primary visual cortex. At that part of the brain, the neurons located at certain columns, respond to particular directions. This orientation changes in a systematic way as one moves across the cortical surface. In our case, the signals from the above-mentioned array are analyzed and information concerning orientation and size of a particular line is obtained. With this system, the Muelle-Lyer illusion has been studied and some rules to interpret why equal length objects give rise to different interpretations are presented.

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On the basis of optical characterization experiments and an eight band kp model, we have studied the effect of Sb incorporation on the electronic structure of InAs quantum dots (QDs). We have found that Sb incorporation in InAs QDs shifts the hole wave function to the center of the QD from the edges of the QD where it is otherwise pinned down by the effects of shear stress. The observed changes in the ground-state energy cannot merely be explained by a composition change upon Sb exposure but can be accounted for when the change in lateral size is taken into consideration. The Sb distribution inside the QDs produces distinctive changes in the density of states, particularly, in the separation between excitation shells. We find a 50% increase in the thermal escape activation energy compared with reference InAs quantum dots as well as an increment of the fundamental transition decay time with Sb incorporation. Furthermore, we find that Sb incorporation into quantum dots is strongly nonlinear with coverage, saturating at low doses. This suggests the existence of a solubility limit of the Sb incorporation into the quantum dots during growth.

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The study of granular systems is of great interest to many fields of science and technology. The packing of particles affects to the physical properties of the granular system. In particular, the crucial influence of particle size distribution (PSD) on the random packing structure increase the interest in relating both, either theoretically or by computational methods. A packing computational method is developed in order to estimate the void fraction corresponding to a fractal-like particle size distribution.

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The study of soil structure, i.e., the pores, is of vital importance in different fields of science and technology. Total pore volume (porosity), pore surface, pore connectivity and pore size distribution are some (probably the most important) of the geometric measurements of pore space. The technology of X-ray computed tomography allows us to obtain 3D images of the inside of a soil sample enabling study of the pores without disturbing the samples. In this work we performed a set of geometrical measures, some of them from mathematical morphology, to assess and quantify any possible difference that tillage may have caused on the soil. We compared samples from tilled soil with samples from a soil with natural vegetation taken in a very close area. Our results show that the main differences between these two groups of samples are total surface area and pore connectivity per unit pore volume.

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A 2D computer simulation method of random packings is applied to sets of particles generated by a self-similar uniparametric model for particle size distributions (PSDs) in granular media. The parameter p which controls the model is the proportion of mass of particles corresponding to the left half of the normalized size interval [0,1]. First the influence on the total porosity of the parameter p is analyzed and interpreted. It is shown that such parameter, and the fractal exponent of the associated power scaling, are efficient packing parameters, but this last one is not in the way predicted in a former published work addressing an analogous research in artificial granular materials. The total porosity reaches the minimum value for p = 0.6. Limited information on the pore size distribution is obtained from the packing simulations and by means of morphological analysis methods. Results show that the range of pore sizes increases for decreasing values of p showing also different shape in the volume pore size distribution. Further research including simulations with a greater number of particles and image resolution are required to obtain finer results on the hierarchical structure of pore space.

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The study of granular systems is of great interest to many fields of science and technology. The packing of particles affects to the physical properties of the granular system. In particular, the crucial influence of particle size distribution (PSD) on the random packing structure increase the interest in relating both, either theoretically or by computational methods. A packing computational method is developed in order to estimate the void fraction corresponding to a fractal-like particle size distribution.

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The purpose of this study was to compare a number of state-of-the-art methods in airborne laser scan- ning (ALS) remote sensing with regards to their capacity to describe tree size inequality and other indi- cators related to forest structure. The indicators chosen were based on the analysis of the Lorenz curve: Gini coefficient ( GC ), Lorenz asymmetry ( LA ), the proportions of basal area ( BALM ) and stem density ( NSLM ) stocked above the mean quadratic diameter. Each method belonged to one of these estimation strategies: (A) estimating indicators directly; (B) estimating the whole Lorenz curve; or (C) estimating a complete tree list. Across these strategies, the most popular statistical methods for area-based approach (ABA) were used: regression, random forest (RF), and nearest neighbour imputation. The latter included distance metrics based on either RF (NN–RF) or most similar neighbour (MSN). In the case of tree list esti- mation, methods based on individual tree detection (ITD) and semi-ITD, both combined with MSN impu- tation, were also studied. The most accurate method was direct estimation by best subset regression, which obtained the lowest cross-validated coefficients of variation of their root mean squared error CV(RMSE) for most indicators: GC (16.80%), LA (8.76%), BALM (8.80%) and NSLM (14.60%). Similar figures [CV(RMSE) 16.09%, 10.49%, 10.93% and 14.07%, respectively] were obtained by MSN imputation of tree lists by ABA, a method that also showed a number of additional advantages, such as better distributing the residual variance along the predictive range. In light of our results, ITD approaches may be clearly inferior to ABA with regards to describing the structural properties related to tree size inequality in for- ested areas.

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This letter presents a novel temperature sensor, which consists of an interdigitated comb electrode structure with a micrometric-scale size, nanometric metallic layer, and nematic liquid crystal (NLC) film. This sensor exploits the permittivity dependence of the NLC with temperature and principle of electrical conductivity above the percolation threshold in thin film metallic layers. The latter has been demonstrated to increase the temperature sensitivity considerably. The high impedance input reduces the power dissipation, and the high enough voltage output makes it easy to measure the output signal with high precision. The operation principle and fabrication process as well as the characterization of the temperature sensor are presented. Experimental results show that the device offers a sensitivity of 9 mV/°C and is dependent on the applied voltage. This is six times greater than the same structure without the use of a nanometric layer.

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Galileo postulated the existence of an insurmountable size for stone columns bearing a useful load as the size for which the structure is only able to resist its self-weight. Herein a method for the determination of the unsurmountable size for truss-like structures is shown, given the form of these structures and the ratio between the allowable stress and the specific weight of the material (the material structural scope). Three types of bars are considered: straight bars, with solid and hollow rectangular cross-section, and catenary bars with circular cross-section —a limit and theoretical case for estimating a meaningful upper bound of the structural scope—. An approximate rule to estimate the structural efficiency —here named GA rule— is shown, and is compared with numerical solutions using the proposed method.

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El estudio de la estructura del suelo es de vital importancia en diferentes campos de la ciencia y la tecnología. La estructura del suelo controla procesos físicos y biológicos importantes en los sistemas suelo-planta-microorganismos. Estos procesos están dominados por la geometría de la estructura del suelo, y una caracterización cuantitativa de la heterogeneidad de la geometría del espacio poroso es beneficiosa para la predicción de propiedades físicas del suelo. La tecnología de la tomografía computerizada de rayos-X (CT) nos permite obtener imágenes digitales tridimensionales del interior de una muestra de suelo, proporcionando información de la geometría de los poros del suelo y permitiendo el estudio de los poros sin destruir las muestras. Las técnicas de la geometría fractal y de la morfología matemática se han propuesto como una poderosa herramienta para analizar y cuantificar características geométricas. Las dimensiones fractales del espacio poroso, de la interfaz poro-sólido y de la distribución de tamaños de poros son indicadores de la complejidad de la estructura del suelo. Los funcionales de Minkowski y las funciones morfológicas proporcionan medios para medir características geométricas fundamentales de los objetos geométricos tridimensionales. Esto es, volumen, superficie, curvatura media de la superficie y conectividad. Las características del suelo como la distribución de tamaños de poros, el volumen del espacio poroso o la superficie poro-solido pueden ser alteradas por diferentes practicas de manejo de suelo. En este trabajo analizamos imágenes tomográficas de muestras de suelo de dos zonas cercanas con practicas de manejo diferentes. Obtenemos un conjunto de medidas geométricas, para evaluar y cuantificar posibles diferencias que el laboreo pueda haber causado en el suelo. ABSTRACT The study of soil structure is of vital importance in different fields of science and technology. Soil structure controls important physical and biological processes in soil-plant-microbial systems. Those processes are dominated by the geometry of soil pore structure, and a quantitative characterization of the spatial heterogeneity of the pore space geometry is beneficial for prediction of soil physical properties. The technology of X-ray computed tomography (CT) allows us to obtain three-dimensional digital images of the inside of a soil sample providing information on soil pore geometry and enabling the study of the pores without disturbing the samples. Fractal geometry and mathematical morphological techniques have been proposed as powerful tools to analyze and quantify geometrical features. Fractal dimensions of pore space, pore-solid interface and pore size distribution are indicators of soil structure complexity. Minkowski functionals and morphological functions provide means to measure fundamental geometrical features of three-dimensional geometrical objects, that is, volume, boundary surface, mean boundary surface curvature, and connectivity. Soil features such as pore-size distribution, pore space volume or pore-solid surface can be altered by different soil management practices. In this work we analyze CT images of soil samples from two nearby areas with contrasting management practices. We performed a set of geometrical measures, some of them from mathematical morphology, to assess and quantify any possible difference that tillage may have caused on the soil.

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Quercus pyrenaica es una especie rebrotadora de raíz intensa e históricamente aprovechada en monte bajo para la obtención de leñas, carbón y pastos. Debido al éxodo rural y a la aparición de nuevas fuentes energéticas, este aprovechamiento fue abandonado en la década de 1970. Desde entonces, las bajas producciones de madera y bellota y el puntisecado de los pies evidencian el generalizado estancamiento de estas masas. Uno de los mayores retos actuales de la selvicultura en el ámbito mediterráneo es encontrar usos alternativos para estos montes abandonados, siendo la conversión a monte alto una de las alternativas preferidas. Se han realizado resalveos de conversión, sin embrago, éstos se aplican sin un conocimiento integral de las causas de la degradación. En esta tesis doctoral, estudiamos un hipotético desequilibrio entre la parte radical y la parte aérea (R:S) de las cepas de rebollo como causa subyacente de su decaimiento. En una parcela experimental, aprovechada al menos desde el siglo XII, se realizaron análisis genéticos a priori para elucidar la estructura genética del rodal, y así estudiar la influencia del tamaño clonal en el funcionamiento de las cepas. Las cepas de mayor tamaño presentaron un menor crecimiento diametral de sus pies, así como mayores tasas de respiración radical, estimadas a partir de flujos internos de CO2 a través del xilema (FT) y de los flujos de CO2 del suelo. Estos resultados sugieren que el desequilibrio R:S aumenta con el tamaño clonal, dado que la eliminación periódica de órganos aéreos, al mismo tiempo que las raíces permanecen intactas, da lugar a un gran desarrollo del sistema radical que consume gran parte de los carbohidratos no estructurales (NSC) en respiración de mantenimiento, comprometiendo así el desarrollo de órganos aéreos. Se excavaron y pesaron dos cepas compuestas por cuatro y ocho pies, las cuales mostraron ratios R:S (0.5 y 1, respectivamente) superiores a los registrados en pies de origen sexual. Al igual que en otras especies rebrotadoras de raíz, se observaron altas concentraciones de NSC en las raíces (> 20% en primavera) y una gran proporción de albura en el sistema radical (52%) que alberga una notable reserva de NSC (87 kg en la cepa de mayor tamaño). En el sistema radical de dicha cepa, estimada mediante dataciones radiocarbónicas en 550 años de edad, se contaron 248 uniones radicales. La persistencia de sistemas radicales grandes, viejos, y altamente interconectados sugiere que la gran cantidad de recursos almacenados y consumidos en las raíces compensan un pobre desarrollo aéreo con una alta resiliencia vegetativa. Para un mejor entendimiento de los balances de carbono y del agotamiento de NSC en las cepas de rebollo, se midieron los flujos internos y externos de CO2 en troncos y los flujos de CO2 del suelo, y se estimó la respiración de órganos aéreos (RS) y subterráneos (RR). Estacionalmente, RS y RR reflejaron las dinámicas de flujo de savia y de crecimiento del tronco, y estuvieron determinadas principalmente por los flujos externos de CO2, dada la escasa contribución de FT a RS y RR (< 10% y < 2%, respectivamente). En una escala circadiana, la contribución de FT a RS aumentó hasta un 25% en momentos de alta transpiración. Las bajas concentraciones de CO2 en el xilema ([CO2] hasta un 0.11%) determinaron comparativamente unos bajos FT, probablemente causados por una limitada respiración del xilema y una baja resistencia a la difusión radial del CO2 impuestos por la sequía estival. Los pulsos de [CO2] observados tras las primeras lluvias de otoño apoyan esta idea. A lo largo del periodo vegetativo, el flujo medio de CO2 procedente del suelo (39 mol CO2 day-1) fue el mayor flujo respiratorio, tres y cuatro veces superior a RS (12 mol CO2 day-1) y RR (8-9 mol CO2 day-1), respectivamente. Ratios RR/RS menores que la unidad evidencian un importante peso de la respiración aérea como sumidero de carbono adicional. Finalmente, se ensayó el zanjado de raíces y el anillamiento de troncos como tratamientos selvícolas alternativos con el objetivo de aumentar las reservas de NSC en los troncos de las cepas. Los resultados preliminares desaconsejan el zanjado de raíces por el alto coste derivado posiblemente de la cicatrización de las heridas. El anillado de troncos imposibilitó el transporte de NSC a las raíces y aumentó la concentración de almidón por encima de la zona anillada, mientras que sistema radical se mantiene por los pies no anillados de la cepa. Son necesarias más mediciones y datos adicionales para comprobar el mantenimiento de esta respuesta positiva a largo plazo. Para concluir, destacamos la necesidad de estudios multidisciplinares que permitan una comprensión integral de la degradación de los rebollares ibéricos para poder aplicar a posteriori una gestión adecuada en estos montes bajos abandonados. ABSTRACT Quercus pyrenaica is a vigorous root-resprouting species intensively and historically coppiced for firewood, charcoal and woody pastures. Due to the rural exodus and the appearance of new energy sources, coppicing was abandoned towards 1970. Since then, tree overaging has resulted in stand stagnation displayed by slow stem growth, branch dieback, and scarce acorn production. The urgent need to find new alternative uses for abandoned coppices is recognized as one of the biggest challenges which currently faces Mediterranean silviculture; conversion into high forest by thinning is one of the preferred alternatives. For this aim, thinning has been broadly applied and seldom tested, although without a comprehensive understanding of the causes of stand stagnation. In this PhD study, we test the hypothesis of an imbalance between above- and below-ground organs, result of long term coppicing, as the underlying cause of Q. pyrenaica decay. In an experimental plot coppiced since at least the 12th century, genetic analyses were performed a priori to elucidate inconspicuous clonal structure of Q. pyrenaica to evaluate how clonal size affects the functioning of these multi-stemmed trees. Clonal size negatively affected diametric stem growth, whereas root respiration rates, measured by internal fluxes of CO2 through xylem (FT) and soil CO2 efflux, increased with clonal size. These results suggest root-to-shoot (R:S) imbalance intensifying with clonal size: periodic removal of aboveground organs whilst belowground organs remain undisturbed may have led to massive root systems which consume a great proportion of non-structural carbohydrates (NSC) for maintenance respiration, thus constraining aboveground performance. Furthermore, excavation of two multi-stemmed trees, composed by four and eight stems, revealed R:S ratios (0.5 and 1, respectively) greater than those reported for sexually regenerated trees. Moreover, as similarly observed in several root-resprouting species, NSC allocation to roots was favored ([NSC] > 20% in spring): a large proportion of sapwood maintained throughout the root system (52%) stored a remarkable NSC pool of 87 kg in the case of the largest clone. In this root system of the eight-stemmed tree, 248 root connections were counted and, by radiocarbon dating, its age was estimated to be 550-years-old. Persistence of massive, old and highly interconnected root systems suggests that enhanced belowground NSC storage and consumption reflects a trade-off between vegetative resilience and aboveground development. For a better understanding of tree carbon budget and the potential role of carbon starvation in Q. pyrenaica decay, internal and external stem CO2 fluxes and soil CO2 effluxes were monitored to evaluate respiratory costs above- and below-ground. On a seasonal scale, stem and root respiration (RS and RR) mirrored sap flow and stem growth dynamics. Respiration was determined to the greatest extent by external fluxes of CO2 to the atmosphere or soil, since FT accounted for a low proportion of RS and RR (< 10% and < 2%, respectively). On a diel scale, the contribution of FT to RS increased up to 25% at high transpiration rates. Comparatively low FT was determined by the low concentration of xylem CO2 registered ([CO2] as low as 0.11%), likely as a consequence of constrained xylem respiration and reduced resistance to CO2 radial diffusion imposed by summer drought. Xylem [CO2] pulses following first autumn rains support this idea. Averaged over the growing season, soil CO2 efflux was the greatest respiratory flux (39 mol CO2 day-1), three and four times greater than RS (12 mol CO2 day-1) and RR (8-9 mol CO2 day-1), respectively. Ratios of RR/RS below one evidence an additional and important weight of aboveground respiration as a tree carbon sink. Finally, root trenching and stem girdling were tested as complimentary treatments to thinning as a means to improve carbon reserves in stems of clonal trees. Preliminary results discouraged root trenching due to the high cost likely incurred for wound closure. Stem girdling successfully blocked NSC translocation downward, increasing starch concentrations above the girdled zone whilst the root system is fed by non-girdled stems within the clone. Further measurements and ancillary data are necessary to verify that this positive effect hold over time. To conclude, the need of multidisciplinary approaches for an integrative understanding on the functioning of abandoned Q pyrenaica coppices is highlighted for an appropriate management of these stands.

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Existe normalmente el propósito de obtener la mejor solución posible cuando se plantea un problema estructural, entendiendo como mejor la solución que cumpliendo los requisitos estructurales, de uso, etc., tiene un coste físico menor. En una primera aproximación se puede representar el coste físico por medio del peso propio de la estructura, lo que permite plantear la búsqueda de la mejor solución como la de menor peso. Desde un punto de vista práctico, la obtención de buenas soluciones—es decir, soluciones cuyo coste sea solo ligeramente mayor que el de la mejor solución— es una tarea tan importante como la obtención de óptimos absolutos, algo en general difícilmente abordable. Para disponer de una medida de la eficiencia que haga posible la comparación entre soluciones se propone la siguiente definición de rendimiento estructural: la razón entre la carga útil que hay que soportar y la carga total que hay que contabilizar (la suma de la carga útil y el peso propio). La forma estructural puede considerarse compuesta por cuatro conceptos, que junto con el material, definen una estructura: tamaño, esquema, proporción, y grueso.Galileo (1638) propuso la existencia de un tamaño insuperable para cada problema estructural— el tamaño para el que el peso propio agota una estructura para un esquema y proporción dados—. Dicho tamaño, o alcance estructural, será distinto para cada material utilizado; la única información necesaria del material para su determinación es la razón entre su resistencia y su peso especifico, una magnitud a la que denominamos alcance del material. En estructuras de tamaño muy pequeño en relación con su alcance estructural la anterior definición de rendimiento es inútil. En este caso —estructuras de “talla nula” en las que el peso propio es despreciable frente a la carga útil— se propone como medida del coste la magnitud adimensional que denominamos número de Michell, que se deriva de la “cantidad” introducida por A. G. M. Michell en su artículo seminal de 1904, desarrollado a partir de un lema de J. C. Maxwell de 1870. A finales del siglo pasado, R. Aroca combino las teorías de Galileo y de Maxwell y Michell, proponiendo una regla de diseño de fácil aplicación (regla GA), que permite la estimación del alcance y del rendimiento de una forma estructural. En el presente trabajo se estudia la eficiencia de estructuras trianguladas en problemas estructurales de flexión, teniendo en cuenta la influencia del tamaño. Por un lado, en el caso de estructuras de tamaño nulo se exploran esquemas cercanos al optimo mediante diversos métodos de minoración, con el objetivo de obtener formas cuyo coste (medido con su numero deMichell) sea muy próximo al del optimo absoluto pero obteniendo una reducción importante de su complejidad. Por otro lado, se presenta un método para determinar el alcance estructural de estructuras trianguladas (teniendo en cuenta el efecto local de las flexiones en los elementos de dichas estructuras), comparando su resultado con el obtenido al aplicar la regla GA, mostrando las condiciones en las que es de aplicación. Por último se identifican las líneas de investigación futura: la medida de la complejidad; la contabilidad del coste de las cimentaciones y la extensión de los métodos de minoración cuando se tiene en cuenta el peso propio. ABSTRACT When a structural problem is posed, the intention is usually to obtain the best solution, understanding this as the solution that fulfilling the different requirements: structural, use, etc., has the lowest physical cost. In a first approximation, the physical cost can be represented by the self-weight of the structure; this allows to consider the search of the best solution as the one with the lowest self-weight. But, from a practical point of view, obtaining good solutions—i.e. solutions with higher although comparable physical cost than the optimum— can be as important as finding the optimal ones, because this is, generally, a not affordable task. In order to have a measure of the efficiency that allows the comparison between different solutions, a definition of structural efficiency is proposed: the ratio between the useful load and the total load —i.e. the useful load plus the self-weight resulting of the structural sizing—. The structural form can be considered to be formed by four concepts, which together with its material, completely define a particular structure. These are: Size, Schema, Slenderness or Proportion, and Thickness. Galileo (1638) postulated the existence of an insurmountable size for structural problems—the size for which a structure with a given schema and a given slenderness, is only able to resist its self-weight—. Such size, or structural scope will be different for every different used material; the only needed information about the material to determine such size is the ratio between its allowable stress and its specific weight: a characteristic length that we name material structural scope. The definition of efficiency given above is not useful for structures that have a small size in comparison with the insurmountable size. In this case—structures with null size, inwhich the self-weight is negligible in comparisonwith the useful load—we use as measure of the cost the dimensionless magnitude that we call Michell’s number, an amount derived from the “quantity” introduced by A. G. M. Michell in his seminal article published in 1904, developed out of a result from J. C.Maxwell of 1870. R. Aroca joined the theories of Galileo and the theories of Maxwell and Michell, obtaining some design rules of direct application (that we denominate “GA rule”), that allow the estimation of the structural scope and the efficiency of a structural schema. In this work the efficiency of truss-like structures resolving bending problems is studied, taking into consideration the influence of the size. On the one hand, in the case of structures with null size, near-optimal layouts are explored using several minimization methods, in order to obtain forms with cost near to the absolute optimum but with a significant reduction of the complexity. On the other hand, a method for the determination of the insurmountable size for truss-like structures is shown, having into account local bending effects. The results are checked with the GA rule, showing the conditions in which it is applicable. Finally, some directions for future research are proposed: the measure of the complexity, the cost of foundations and the extension of optimization methods having into account the self-weight.

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Nowadays, translating information about hydrologic and soil properties and processes across scales has emerged as a major theme in soil science and hydrology, and suitable theories for upscaling or downscaling hydrologic and soil information are being looked forward. The recognition of low-order catchments as self-organized systems suggests the existence of a great amount of links at different scales between their elements. The objective of this work was to research in areas of homogeneous bedrock material, the relationship between the hierarchical structure of the drainage networks at hillslope scale and the heterogeneity of the particle-size distribution at pedon scale. One of the most innovative elements in this work is the choice of the parameters to quantify the organization level of the studied features. The fractal dimension has been selected to measure the hierarchical structure of the drainage networks, while the Balanced Entropy Index (BEI) has been the chosen parameter to quantify the heterogeneity of the particle-size distribution from textural data. These parameters have made it possible to establish quantifiable relationships between two features attached to different steps in the scale range. Results suggest that the bedrock lithology of the landscape constrains the architecture of the drainage networks developed on it and the particle soil distribution resulting in the fragmentation processes.

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Nowadays, translating information about hydrologic and soil properties and processes across scales has emerged as a major theme in soil science and hydrology, and suitable theories for upscaling or downscaling hydrologic and soil information are being looked forward. The recognition of low-order catchments as self-organized systems suggests the existence of a great amount of links at different scales between their elements. The objective of this work was to research in areas of homogeneous bedrock material, the relationship between the hierarchical structure of the drainage networks at hillslope scale and the heterogeneity of the particle-size distribution at pedon scale. One of the most innovative elements in this work is the choice of the parameters to quantify the organization level of the studied features. The fractal dimension has been selected to measure the hierarchical structure of the drainage networks, while the Balanced Entropy Index (BEI) has been the chosen parameter to quantify the heterogeneity of the particle-size distribution from textural data. These parameters have made it possible to establish quantifiable relationships between two features attached to different steps in the scale range. Results suggest that the bedrock lithology of the landscape constrains the architecture of the drainage networks developed on it and the particle soil distribution resulting in the fragmentation processes.

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The RNA phage Qβ requires for the replication of its genome an RNA binding protein called Qβ host factor or Hfq protein. Our previous results suggested that this protein mediates the access of replicase to the 3′-end of the Qβ plus strand RNA. Here we report the results of an evolutionary experiment in which phage Qβ was adapted to an Escherichia coli Q13 host strain with an inactivated host factor (hfq) gene. This strain initially produced phage at a titer ≈10,000-fold lower than the wild-type strain and with minute plaque morphology, but after 12 growth cycles, phage titer and plaque size had evolved to levels near those of the wild-type host. RNAs isolated from adapted Qβ mutants were efficient templates for replicase without host factor in vitro. Electron microscopy showed that mutant RNAs, in contrast to wild-type RNA, efficiently interacted with replicase at the 3′-end in the absence of host factor. The same set of four mutations in the 3′-terminal third of the genome was found in several independently evolved phage clones. One mutation disrupts the base pairing of the 3′-terminal CCCoh sequence, suggesting that the host factor stimulates activity of the wild-type RNA template by melting out its 3′-end.