Changes in spatial point patterns of pioneer woody plants across a large tropical landslide

We assessed whether the relative importance of positive and negative interactions in early successional communities varied across a large landslide on Casita Volcano (Nicaragua). We tested several hypotheses concerning the signatures of these processes in the spatial patterns of woody pioneer plants, as well as those of mortality and recruitment events, in several zones of the landslide differing in substrate stability and fertility, over a period of two years (2001 and 2002). We identified all woody individuals with a diameter >1 cm and mapped them in 28 plots measuring 10 × 10-m. On these maps, we performed a spatial point pattern analysis using univariate and bivariate pair-correlation functions; g (r) and g12 (r), and pairwise differences of univariate and bivariate functions. Spatial signatures of positive and negative interactions among woody plants were more prevalent in the most and least stressful zones of the landslide, respectively. Natural and human-induced disturbances such as the occurrence of fire, removal of newly colonizing plants through erosion and clearcutting of pioneer trees were also identified as potentially important pattern-creating processes. These results are in agreement with the stress-gradient hypothesis, which states that the relative importance of facilitation and competition varies inversely across gradients of abiotic stress. Our findings also indicate that the assembly of early successional plant communities in large heterogeneous landslides might be driven by a much larger array of processes than previously thought.

Three-dimensional distribution of cortical synapses: a replicated point pattern-based analysis

The biggest problem when analyzing the brain is that its synaptic connections are extremely complex. Generally, the billions of neurons making up the brain exchange information through two types of highly specialized structures: chemical synapses (the vast majority) and so-called gap junctions (a substrate of one class of electrical synapse). Here we are interested in exploring the three-dimensional spatial distribution of chemical synapses in the cerebral cortex. Recent research has showed that the three-dimensional spatial distribution of synapses in layer III of the neocortex can be modeled by a random sequential adsorption (RSA) point process, i.e., synapses are distributed in space almost randomly, with the only constraint that they cannot overlap. In this study we hypothesize that RSA processes can also explain the distribution of synapses in all cortical layers. We also investigate whether there are differences in both the synaptic density and spatial distribution of synapses between layers. Using combined focused ion beam milling and scanning electron microscopy (FIB/SEM), we obtained three-dimensional samples from the six layers of the rat somatosensory cortex and identified and reconstructed the synaptic junctions. A total volume of tissue of approximately 4500μm3 and around 4000 synapses from three different animals were analyzed. Different samples, layers and/or animals were aggregated and compared using RSA replicated spatial point processes. The results showed no significant differences in the synaptic distribution across the different rats used in the study. We found that RSA processes described the spatial distribution of synapses in all samples of each layer. We also found that the synaptic distribution in layers II to VI conforms to a common underlying RSA process with different densities per layer. Interestingly, the results showed that synapses in layer I had a slightly different spatial distribution from the other layers.

Estudio de los factores que condicionan, a escala local, la presencia de Epidendrum rhopalostele (Orchidaceae): Implicaciones para su conservación

El análisis de los factores que determinan el establecimiento y supervivencia de orquídeas epífitas, incluyen: a) las condiciones microambientales de los bosques que las mantienen, b) preferencias por las características de los hospederos donde crecen, c) limitación en la dispersión de semillas, d) interacciones planta-planta, y e) asociaciones micorrízicas para la germinación y resultan esenciales para el desarrollo de estrategias para la conservación y manejo de este grupo de plantas. Este trabajo ha evaluado la importancia de estos factores en Epidendrum rhopalostele, orquídea epífita del bosque de niebla montano, a través de los análisis de los patrones espaciales de los árboles que la portan y de la propia orquídea, a escala de población, estudios de asociación y métodos moleculares. Estos últimos han consistido en el uso de marcadores AFLP para el análisis de la estructura genética de la orquídea y en la secuenciación-clonación de la región ITS para la identificación de los hongos micorrízicos asociados. El objetivo de esta tesis es, por tanto, una mejor comprensión de los factores que condicionan la presencia de orquídeas epífitas en los remanentes de bosque de niebla montano y una evaluación de las implicaciones para la conservación y mantenimiento de sus hábitats y la permanencia de sus poblaciones. El estudio fue realizado en un fragmento de bosque de niebla montano de sucesión secundaria situado al este de la Cordillera Real, en los Andes del sur de Ecuador, a 2250 m.s.n.m y caracterizado por una pendiente marcada, temperatura media anual de 20.8°C y precipitación anual de 2193 mm. En este fragmento se mapearon, identificaron y caracterizaron todos los árboles presentes con DBH > 1 cm y todos los individuos de Epidendrum rhopalostele. Así mismo se tomaron muestras de hoja para obtener ADN de todas las orquídeas registradas y muestras de raíces de individuos con flor de E. rhopalostele, uno por cada forófito, para el análisis filogenético de micorrizas. Análisis espaciales de patrones de puntos basados en la K de Ripley y la distancia al vecino más cercano fueron usados para los árboles, forófitos y la población de E. rhopalostele. Se observó que la distribución espacial de árboles y forófitos de E. rhopalostele no es aleatoria, ya que se ajusta a un proceso agregado de Poisson. De ahí se infiere una limitación en la dispersión de las semillas en el fragmento estudiado y en el establecimiento de la orquídea. El patrón de distribución de la población de E. rhopalostele en el fragmento muestra un agrupamiento a pequeña escala sugiriendo una preferencia por micro-sitios para el establecimiento de la orquídea con un kernel de dispersión de las semillas estimado de 0.4 m. Las características preferentes del micro-sitio como tipos de árboles (Clusia alata y árboles muertos), tolerancia a la sombra, corteza rugosa, distribución en los dos primeros metros sugieren una tendencia a distribuirse en el sotobosque. La existencia de una segregación espacial entre adultos y juveniles sugiere una competencia por recursos limitados condicionada por la preferencia de micro-sitio. La estructura genética de la población de E. rhopalostele analizada a través de Structure y PCoA evidencia la presencia de dos grupos genéticos coexistiendo en el fragmento y en los mismos forófitos, posiblemente por eventos de hibridización entre especies de Epidendrum simpátricas. Los resultados del análisis de autocorrelación espacial efectuados en GenAlex confirman una estructura genético-espacial a pequeña escala que es compatible con un mecanismo de dispersión de semillas a corta distancia ocasionada por gravedad o pequeñas escorrentías, frente a la dispersión a larga distancia promovida por el viento generalmente atribuida a las orquídeas. Para la identificación de los micobiontes se amplificó la región ITS1-5.8S-ITS2, y 47 secuencias fueron usadas para el análisis filogenético basado en neighborjoining, análisis bayesiano y máximum-likelihood que determinó que Epidendrum rhopalostele establece asociaciones micorrízicas con al menos dos especies diferentes de Tulasnella. Se registraron plantas que estaban asociadas con los dos clados de hongos encontrados, sugiriendo ausencia de limitación en la distribución del hongo. Con relación a las implicaciones para la conservación in situ resultado de este trabajo se recomienda la preservación de todo el fragmento de bosque así como de las interacciones existentes (polinizadores, micorrizas) a fin de conservar la diversidad genética de esta orquídea epífita. Si fuere necesaria una reintroducción se deben contemplar distancias entre los individuos en cada forófito dentro de un rango de 0.4 m. Para promover el reclutamiento y regeneración de E. rhopalostele, se recomienda que los forófitos correspondan preferentemente a árboles muertos o caídos y a especies, como Clusia alata, que posean además corteza rugosa, sean tolerantes a la sombra, y en el área del sotobosque con menor luminosidad. Además es conveniente que las orquídeas en su distribución vertical estén ubicadas en los primeros metros. En conclusión, la limitación en la dispersión, las características del micro-sitio, las interacciones intraespecíficas y con especies congenéricas simpátricas y las preferencias micorrízicas condicionan la presencia de esta orquídea epífita en este tipo de bosque. ABSTRACT The analysis of factors that determine the establishment and survival of epiphytic depends on factors such as a) microenvironmental conditions of forest, b) preference for host characteristics where orchids grow, c) seed dispersal limitation, d) plant-plant interaction, e) priority mycorrhizal associations for germination, are essential for the development of strategies for management and conservation. This work evaluated the importance of these factors in Epidendrum rhopalostele, an epiphytic orchid of montane cloud forest through the analysis of spatial patterns of host trees and the orchid, in a more specific scale, with association studies and molecular methods, including AFLPs for orchid population genetic structure and the sequencing of the ITS region for associated mycorrhizal fungi. The aim of this thesis is to understand the factors that condition the presence of epiphytic orchids in the remnants of montane cloud forest and to assess the implications for the conservation and preservation of their habitats and the persistence of the orchid populations. The study was carried out in a fragment of montane cloud forest of secondary succession on the eastern slope of Cordillera Real in the Andes of southern Ecuador, located at 2250 m a.s.l. characterized by a steep slope, mean annual temperature of 20.8°C and annual precipitation of 2193 mm. All trees with DBH > 1 cm were mapped, characterized and identified. All E. rhopalostele individuals present were counted, marked, characterized and mapped. Leaf samples of all orchid individuals were collected for DNA analysis. Root samples of flowering E. rhopalostele individuals were collected for phylogenetic analysis of mycorrhizae, one per phorophyte. Spatial point pattern analysis based on Ripley`s K function and nearest neighbor function was used for trees, phorophytes and orchid population. We observed that spatial distribution of trees and phorophytes is not random, as it adjusts to a Poisson cluster process. This suggests a limitation for seed dispersal in the study fragment that is affecting orchid establishment. Furthermore, the small-scale spatial pattern of E. rhopalostele evidences a clustering that suggests a microsite preference for orchid establishment with a dispersal kernel of 0.4 m. Microsite features such as types of trees (dead trees or Clusia alata), shade tolerance trees, rough bark, distribution in the first meters suggest a tendency to prefer the understory for their establishment. Regarding plant-plant interaction a spatial segregation between adults and juveniles was present suggesting competition for limited resources conditioned for a microsite preference. Analysis of genetic structure of E. rhopalostele population through Structure and PCoA shows two genetic groups coexisting in this fragment and in the same phorophyte, possibly as a result of hybridization between sympatric species of Epidendrum. Our results of spatial autocorrelation analysis develop in GenAlex confirm a small-scale spatial-genetic structure within the genetic groups that is compatible with a short-distance dispersal mechanism caused by gravity or water run-off, instead of the long-distance seed dispersal promoted by wind generally attributed to orchids. For mycobionts identification ITS1-5.8S-ITS2 rDNA region was amplified. Phylogenetic analysis was performed with neighborjoining, Bayesian likelihood and maximum-likelihood for 47 sequences yielded two Tulasnella clades. This orchid establishes mycorrhizal associations with at least two different Tulasnella species. In some cases both fungi clades were present in same root, suggesting no limitation in fungal distribution. Concerning the implications for in situ conservation resulting from this work, the preservation of all forest fragment and their interactions (pollinators, mycorrhiza) is recommended to conserve the genetic diversity of this species. If a reintroduction were necessary, distances between individuals in each phorophyte within a range of 0.4 m, are recommended. To promote recruitment and regeneration of E. rhopalostele it is recommended that phorophytes correspond to dead or fallen trees or species, such as Clusia alata. Trees that have rough bark and are shade tolerant are also recommended. Furthermore, regarding vertical distribution, it is also convenient that orchids are located in the first meter (in understory, area with less light). In conclusion, limitation on seed dispersal, microsite characteristics, plant-plant interactions or interaction with cogeneric sympatric species and mycorrhizal preferences conditioned the presence of this epiphytic orchid in this fragment forest.

Fine spatial pattern of an epiphytic lichen species is affected by habitat conditions in two forest types in the Iberian Mediterranean region

Persistence and abundance of species is determined by habitat availability and the ability to disperse and colonize habitats at contrasting spatial scales. Favourable habitat fragments are also heterogeneous in quality, providing differing opportunities for establishment and affecting the population dynamics of a species. Based on these principles, we suggest that the presence and abundance of epiphytes may reflect their dispersal ability, which is primarily determined by the spatial structure of host trees, but also by host quality. To our knowledge there has been no explicit test of the importance of host tree spatial pattern for epiphytes in Mediterranean forests. We hypothesized that performance and host occupancy in a favourable habitat depend on the spatial pattern of host trees, because this pattern affects the dispersal ability of each epiphyte and it also determines the availability of suitable sites for establishment. We tested this hypothesis using new point pattern analysis tools and generalized linear mixed models to investigate the spatial distribution and performance of the epiphytic lichen Lobaria pulmonaria, which inhabits two types of host trees (beeches and Iberian oaks). We tested the effects on L. pulmonaria distribution of tree size, spatial configuration, and host tree identity. We built a model including tree size, stand structure, and several neighbourhood predictors to understand the effect of host tree on L. pulmonaria. We also investigated the relative importance of spatial patterning on the presence and abundance of the species, independently of the host tree configuration. L. pulmonaria distribution was highly dependent on habitat quality for successful establishment, i.e., tree species identity, tree diameter, and several forest stand structure surrogates. For beech trees, tree diameter was the main factor influencing presence and cover of the lichen, although larger lichen-colonized trees were located close to focal trees, i.e., young trees. However, oak diameter was not an important factor, suggesting that bark roughness at all diameters favoured lichen establishment. Our results indicate that L. pulmonaria dispersal is not spatially restricted, but it is dependent on habitat quality. Furthermore, new spatial analysis tools suggested that L. pulmonaria cover exhibits a distinct pattern, although the spatial pattern of tree position and size was random.

Fine spatial pattern of an epiphytic lichen species is affected by habitat conditions in two forest types in the Iberian Mediterranean region

Persistence and abundance of species is determined by habitat availability and the ability to disperse and colonize habitats at contrasting spatial scales. Favourable habitat fragments are also heterogeneous in quality, providing differing opportunities for establishment and affecting the population dynamics of a species. Based on these principles, we suggest that the presence and abundance of epiphytes may reflect their dispersal ability, which is primarily determined by the spatial structure of host trees, but also by host quality. To our knowledge there has been no explicit test of the importance of host tree spatial pattern for epiphytes in Mediterranean forests. We hypothesized that performance and host occupancy in a favourable habitat depend on the spatial pattern of host trees, because this pattern affects the dispersal ability of each epiphyte and it also determines the availability of suitable sites for establishment. We tested this hypothesis using new point pattern analysis tools and generalized linear mixed models to investigate the spatial distribution and performance of the epiphytic lichen Lobaria pulmonaria, which inhabits two types of host trees (beeches and Iberian oaks). We tested the effects on L. pulmonaria distribution of tree size, spatial configuration, and host tree identity. We built a model including tree size, stand structure, and several neighbourhood predictors to understand the effect of host tree on L. pulmonaria. We also investigated the relative importance of spatial patterning on the presence and abundance of the species, independently of the host tree configuration. L. pulmonaria distribution was highly dependent on habitat quality for successful establishment, i.e., tree species identity, tree diameter, and several forest stand structure surrogates. For beech trees, tree diameter was the main factor influencing presence and cover of the lichen, although larger lichen-colonized trees were located close to focal trees, i.e., young trees. However, oak diameter was not an important factor, suggesting that bark roughness at all diameters favoured lichen establishment. Our results indicate that L. pulmonaria dispersal is not spatially restricted, but it is dependent on habitat quality. Furthermore, new spatial analysis tools suggested that L. pulmonaria cover exhibits a distinct pattern, although the spatial pattern of tree position and size was random.

Negative density dependence and environmental heterogeneity effects on tree ferns across succession in a tropical montane forest.

Although tree ferns are an important component of temperate and tropical forests, very little is known about their ecology. Their peculiar biology (e.g., dispersal by spores and two-phase life cycle) makes it difficult to extrapolate current knowledge on the ecology of other tree species to tree ferns. In this paper, we studied the effects of negative density dependence (NDD) and environmental heterogeneity on populations of two abundant tree fern species, Cyathea caracasana and Alsophila engelii, and how these effects change across a successional gradient. Species patterns harbor information on processes such as competition that can be easily revealed using point pattern analysis techniques. However, its detection may be difficult due to the confounded effects of habitat heterogeneity. Here, we mapped three forest plots along a successional gradient in the montane forests of Southern Ecuador. We employed homogeneous and inhomogeneous K and pair correlation functions to quantify the change in the spatial pattern of different size classes and a case-control design to study associations between juvenile and adult tree ferns. Using spatial estimates of the biomass of four functional tree types (short- and long-lived pioneer, shade- and partial shade-tolerant) as covariates, we fitted heterogeneous Poisson models to the point pattern of juvenile and adult tree ferns and explored the existence of habitat dependencies on these patterns. Our study revealed NDD effects for C. caracasana and strong environmental filtering underlying the pattern of A. engelii. We found that adult and juvenile populations of both species responded differently to habitat heterogeneity and in most cases this heterogeneity was associated with the spatial distribution of biomass of the four functional tree types. These findings show the effectiveness of factoring out environmental heterogeneity to avoid confounding factors when studying NDD and demonstrate the usefulness of covariate maps derived from mapped communities.

Patrones y procesos espaciales en poblaciones y comunidades vegetales: nuevas herramientas e hipótesis

Generalmente los patrones espaciales de puntos en ecología, se definen en el espacio bi-dimensional, donde cada punto representado por el par ordenado (x,y), resume la ubicación espacial de una planta. La importancia de los patrones espaciales de plantas radica en que proceden como respuesta ante importantes procesos ecológicos asociados a la estructura de una población o comunidad. Tales procesos incluyen fenómenos como la dispersión de semillas, la competencia por recursos, la facilitación, respuesta de las plantas ante algún tipo de estrés, entre otros. En esta tesis se evalúan los factores y potenciales procesos subyacentes, que explican los patrones de distribución espacial de la biodiversidad vegetal en diferentes ecosistemas como bosque mediterráneo, bosque tropical y matorral seco tropical; haciendo uso de nuevas metodologías para comprobar hipótesis relacionadas a los procesos espaciales. En este trabajo se utilizaron dos niveles ecológicos para analizar los procesos espaciales, el nivel de población y el nivel de comunidad, con el fin de evaluar la importancia relativa de las interacciones intraespecíficas e interespecíficas. Me centré en el uso de funciones estadísticas que resumen los patrones de puntos para explorar y hacer inferencias a partir de datos espaciales, empezando con la construcción de un nuevo modelo nulo para inferir variantes del síndrome de dispersión de una planta parásita en España central. Se analizó la dependencia de los patrones espaciales tanto de los hospedantes afectados como de los no-afectados y se observó fuerte dependencia a pequeña y mediana distancia. Se utilizaron dos funciones (kernel) para simular la dispersión de la especie parásita y se identificó consistencia de estos modelos con otros síndromes de dispersión adicionalmente a la autodispersión. Un segundo tema consistió en desarrollar un método ANOVA de dos vías? para patrones de puntos replicados donde el interés se concentró en evaluar la interacción de dos factores. Este método se aplicó a un caso de estudio que consitió en analizar la influencia de la topografía y la altitud sobre el patrón espacial de un arbusto dominante en matorral seco al sur del Ecuador, cuyos datos provienen de patrones de puntos replicados basados en diseño. Partiendo de una metodología desarrollada para procesos uni-factoriales, se construyó el método para procesos bi-factoriales y así poder evaluar el efecto de interacción. Se observó que la topografía por sí sola así como la interacción con la altitud presentaron efecto significativo sobre la formación del patrón espacial. Un tercer tema fue identificar la relación entre el patrón espacial y el síndrome de dispersión de la comunidad vegetal en el bosque tropical de la Isla de Barro Colorado (BCI), Panamá. Muchos estudios se han desarrollado en este bosque tropical y algunos han analizado la relación síndrome-patrón espacial, sin embargo lo novedoso de nuestro estudio es que se evaluaron un conjunto amplio de modelos (114 modelos) basados en procesos que incorporan la limitación de la dispersión y la heterogeneidad ambiental, y evalúan el efecto único y el efecto conjunto, para posteriormente seleccionar el modelo de mejor ajuste para cada especie. Más de la mitad de las especies presentaron patrón espacial consistente con el efecto conjutno de la limitación de la dispersión y heterogeneidad ambiental y el porcentaje restante de especies reveló en forma equitativa el efecto único de la heterogeneidad ambiental y efecto único de limitación de la dispersión. Finalmente, con la misma información del bosque tropical de BCI, y para entender las relaciones que subyacen para mantener el equilibrio de la biodiversidad, se desarrolló un índice de dispersión funcional local a nivel de individuo, que permita relacionar el patrón espacial con cuatro rasgos funcionales clave de las especies. Pese a que muchos estudios realizados involucran esta comunidad con la teoría neutral, se encontró que el ensamble de la comunidad de BCI está afectado por limitaciones de similaridad y de hábitat a diferentes escalas. ABSTRACT Overall the spatial point patterns in ecology are defined in two-dimensional space, where each point denoted by the (x,y) ordered pair, summarizes the spatial location of a plant. The spatial point patterns are essential because they arise in response to important ecological processes, associated with the structure of a population or community. Such processes include phenomena as seed dispersal, competition for resources, facilitation, and plant response to some type of stress, among others. In this thesis, some factors and potential underlying processes were evaluated in order to explain the spatial distribution patterns of plant biodiversity. It was done in different ecosystems such as Mediterranean forest, tropical forest and dry scrubland. For this purpose new methodologies were used to test hypothesis related to spatial processes. Two ecological levels were used to analyze the spatial processes, at population and community levels, in order to assess the relative importance of intraspecific and interspecific interactions. I focused on the use of spatial statistical functions to summarize point patterns to explore and make inferences from spatial data, starting with the construction of a new null model to infer variations about the dispersal syndrome of a parasitic plant in central Spain. Spatial dependence between point patterns in a multivariate point process of affected and unaffected hosts were analyzed and strong dependence was observed at small and medium distance. Two kernel functions were used to simulate the dispersion of parasitic plant and consistency of these models with other syndromes was identified, in addition to ballistic dispersion. A second issue was to analyze altitude and topography effects on the spatial population structure of a dominant shrub in the dry ecosystem in southern Ecuador, whose data come from replicated point patterns design-based. Based on a methodology developed for uni-factorial process, a method for bi-factorial processes was built to assess the interaction effect. The topography alone and interacting with altitude showed significant effect on the spatial pattern of shrub. A third issue was to identify the relationship between the spatial pattern and dispersal syndromes of plant community in the tropical forest of Barro Colorado Island (BCI), Panamá. Several studies have been developed in this tropical forest and some focused on the spatial pattern-syndrome relationship; however the novelty of our study is that a large set of models (114 models) including dispersal limitation and environmental heterogeneity were evaluated, used to identify the only and joint effect to subsequently select the best fit model for each species. Slightly more than fifty percent of the species showed spatial pattern consistent with only the dispersal limitation, and the remaining percentage of species revealed the only effect of environmental heterogeneity and habitat-dispersal limitation joined effect, equitably. Finally, with the same information from the tropical forest of BCI, and to understand the relationships underlying for balance of biodiversity, an index of the local functional dispersion was developed at the individual level, to relate the spatial pattern with four key functional traits of species. Although many studies involve this community with neutral theory, the assembly of the community is affected by similarity and habitat limitations at different scales.

Analysis and design of reconfigurable multimode multiport microstrip antennas for mimo systems and diversity schemes

Las comunicaciones inalámbricas han transformado profundamente la forma en la que la gente se comunica en el día a día y es, sin lugar a dudas, una de las tecnologías de nuestro tiempo que más rápidamente evoluciona. Este rápido crecimiento implica retos enormes en la tecnología subyacente, debido y entre otros motivos, a la gran demanda de capacidad de los nuevos servicios inalámbricos. Los sistemas Multiple Input Multiple Output (MIMO) han despertado mucho interés como medio de mejorar el rendimiento global del sistema, satisfaciendo de este modo y en cierta medida los nuevo requisitos exigidos. De hecho, el papel relevante de esta tecnología en los actuales esfuerzos de estandarización internacionales pone de manifiesto esta utilidad. Los sistemas MIMO sacan provecho de los grados de libertad espaciales, disponibles a través del entorno multitrayecto, para mejorar el rendimiento de la comunicación con una destacable eficiencia espectral. Con el fin de alcanzar esta mejora en el rendimiento, la diversidad espacial y por diagrama han sido empleadas tradicionalmente para reducir la correlación entre los elementos radiantes, ya que una correlación baja es condición necesaria, si bien no suficiente, para dicha mejora. Tomando como referencia, o punto de partida, las técnicas empleadas para obtener diversidad por diagrama, esta tesis doctoral surge de la búsqueda de la obtención de diversidad por diagrama y/o multiplexación espacial a través del comportamiento multimodal de la antena microstrip, proponiendo para ello un modelo cuasi analítico original para el análisis y diseño de antenas microstrip multipuerto, multimodo y reconfigurables. Este novedoso enfoque en este campo, en vez de recurrir a simulaciones de onda completa por medio de herramientas comerciales tal y como se emplea en las publicaciones existentes, reduce significativamente el esfuerzo global de análisis y diseño, en este último caso por medio de guías de diseño generales. Con el fin de lograr el objetivo planteado y después de una revisión de los principales conceptos de los sistemas MIMO que se emplearán más adelante, se fija la atención en encontrar, implementar y verificar la corrección y exactitud de un modelo analítico que sirva de base sobre la cual añadir las mejoras necesarias para obtener las características buscadas del modelo cuasi analítico propuesto. Posteriormente y partiendo del modelo analítico base seleccionado, se exploran en profundidad y en diferentes entornos multitrayecto, las posibilidades en cuanto a rendimiento se refiere de diversidad por diagrama y multiplexación espacial, proporcionadas por el comportamiento multimodal de las antenas parche microstrip sin cargar. Puesto que cada modo de la cavidad tiene su propia frecuencia de resonancia, es necesario encontrar formas de desplazar la frecuencia de resonancia de cada modo empleado para ubicarlas en la misma banda de frecuencia, manteniendo cada modo al mismo tiempo tan independiente como sea posible. Este objetivo puede lograrse cargando adecuadamente la cavidad con cargas reactivas, o alterando la geometría del parche radiante. Por consiguiente, la atención en este punto se fija en el diseño, implementación y verificación de un modelo cuasi analítico para el análisis de antenas parche microstrip multipuerto, multimodo y cargadas que permita llevar a cabo la tarea indicada, el cuál es una de las contribuciones principales de esta tesis doctoral. Finalmente y basándose en el conocimiento adquirido a través del modelo cuasi analítico, se proporcionan y aplican guías generales para el diseño de antenas microstrip multipuerto, multimodo y reconfigurables para sistemas MIMO, con el fin de mejorar su diversidad por diagrama y/o su capacidad por medio del comportamiento multimodal de las antenas parche microstrip. Se debe destacar que el trabajo presentado en esta tesis doctoral ha dado lugar a una publicación en una revista técnica internacional de un alto factor de impacto. De igual manera, el trabajo también ha sido presentado en algunas de las más importantes conferencias internacionales en el ámbito de las antenas ABSTRACT Wireless communications have deeply transformed the way people communicate on daily basis and it is undoubtedly one of the most rapidly evolving technologies of our time. This fast growing behaviour involves huge challenges on the bearing technology, due to and among others reasons, the high demanding capacity of new wireless services. MIMO systems have given rise to considerable interest as a means to enhance the overall system performance, thus satisfying somehow the new demanding requirements. Indeed, the significant role of this technology on current international standardization efforts, highlights this usefulness. MIMO systems make profit from the spatial degrees of freedom available through the multipath scenario to improve the communication performance with a remarkable spectral efficiency. In order to achieve this performance improvement, spatial and pattern diversity have been traditionally used to decrease the correlation between antenna elements, as low correlation is a necessary but not sufficient condition. Taking as a reference, or starting point, the techniques used to achieve pattern diversity, this Philosophiae Doctor (Ph.D.) arises from the pursuit of obtaining pattern diversity and/or spatial multiplexing capabilities through the multimode microstrip behaviour, thus proposing a novel quasi analytical model for the analysis and design of reconfigurable multimode multiport microstrip antennas. This innovative approach on this field, instead of resorting to full-wave simulations through commercial tools as done in the available publications, significantly reduces the overall analysis and design effort, in this last case through comprehensive design guidelines. In order to achieve this goal and after a review of the main concepts of MIMO systems which will be followed used, the spotlight is fixed on finding, implementing and verifying the correctness and accuracy of a base quasi analytical model over which add the necessary enhancements to obtain the sought features of the quasi analytical model proposed. Afterwards and starting from the base quasi analytical model selected, the pattern diversity and spatial multiplexing performance capabilities provided by the multimode behaviour of unloaded microstrip patch antennas under different multipath environments are fully explored. As each cavity mode has its own resonant frequency, it is required to find ways to displace the resonant frequency of each used mode to place them at the same frequency band while keeping each mode as independent as possible. This objective can be accomplished with an appropriate loading of the cavity with reactive loads, or through the alteration of the geometry of the radiation patch. Thus, the focus is set at this point on the design, implementation and verification of a quasi analytical model for the analysis of loaded multimode multiport microstrip patch antennas to carry out the aforementioned task, which is one of the main contributions of this Ph.D. Finally and based on the knowledge acquired through the quasi analytical model, comprehensive guidelines to design reconfigurable multimode MIMO microstrip antennas to improve the spatial multiplexing and/or diversity system performance by means of the multimode microstrip patch antenna behaviour are given and applied. It shall be highlighted that the work presented in this Ph.D. has given rise to a publication in an international technical journal of high impact factor. Moreover, the work has also been presented at some of the most important international conferences in antenna area.

Un entendimiento infraestructural del proyecto arquitectónico

Un entendimiento infraestructural del proyecto arquitectónico Este trabajo de tesis reivindica la pertinencia de la lógica infraestructural como herramienta de proyecto arquitectónico en el territorio urbano contemporáneo, a través del relato del tránsito que se produce entre las décadas de 1960 y 1970 y el final del siglo XX, desde unos sistemas urbanos centrales, vectoriales y ‘modernos’ hacia unos sistemas policéntricos, organizados según modelos de campos y ‘contemporáneos’. Las infraestructuras han sido tradicionalmente objeto de interés para generaciones de arquitectos y urbanistas que, bien desde su condición plástica más pregnante, bien desde una intuición de su capacidad organizativa, han sido fascinados por la irrupción en el territorio de estos elementos generalmente ajenos al proyecto arquitectónico. Los proyectos que durante décadas han explorado la posibilidad de habitar estas grandes estructuras, o de proyectar con su lenguaje y su escala, han resultado en un acercamiento a la capacidad de lo infraestructural como herramienta de proyecto limitada a cuestiones morfotipológicas. Sin embargo, a partir de un estudio de la infraestructura ‘clásica’ como aquella resultado de la manifestación de las redes de abastecimiento y conectividad básicas, se desvela una naturaleza de lo infraestructural que trasciende su expresión formal conocida y explora su capacidad performativa, como sistema de relaciones, como orden topológico, como soporte flexible para la actividad espontánea y no anticipada. A partir del trabajo de diversos autores y de una selección de proyectos de distinta índole –la mayoría no construidos– se muestra cómo la irrupción de una conciencia relacional, así como la problemática moderna de afrontar los procesos de generación de una ‘nueva ciudad para una nueva sociedad’, motivan la búsqueda de sistemas de generación y control de ‘forma colectiva’ adecuados a un modelo urbano contemporáneo. Transitando por los escritos de Fumihiko Maki en Investigations in Collective Form, Reyner Banham en Megastructure, o Shadrach Woods en The Man in the Street durante los 60 y 70 y, más recientemente, de Stan Allen en Points + Lines, Edward Soja en Postmetropolis o Frei Otto en Occupying and Connecting, se traza esta historia semántica de lo infraestructural desde lo tipológico hasta una noción ampliada que resulta pertinente para operar en el territorio contemporáneo postmetropolitano. Para ello el trabajo plantea, por un lado, la descripción del contexto urbano que podemos asumir como contemporáneo, su evolución y sus características principales. Por otro lado, se explora la formación de esta noción de entendimiento infraestructural en las últimas décadas desde enfoques complementarios. Una introducción describe las condiciones de contexto en las que este estudio doctoral adquiere su pertinencia: el tránsito relativamente reciente desde un modelo vectorial y clásico de ocupación del territorio, al modelo contemporáneo disperso y policéntrico propio del modelo de campos. Un bloque central de tres capítulos analiza ‘lo infraestructural’ como expresión de una arquitectura de las relaciones, como soporte para una sociedad en cambio y como lógica proyectual, no ligada a una forma dada. La última parte, a modo de epílogo, describe el territorio urbano contemporáneo, sus cualidades básicas y la capacidad de las lógicas infraestructurales enunciadas en los capítulos anteriores para operar en él. Un último apartado de conclusiones, establece las bases de una línea de investigación abierta acerca de la operatividad actual de estos sistemas en el proyecto de arquitectura. ABSTRACT An infrastructural understanding of architectural design This thesis project claims for the suitability of infrastructural logics as a design tool within contemporary urban territories. This claim is illustrated by means of a report on the transit between the 60–70’s decades and the end of the 20th century, ranging from vector, central, ‘modern’ urban systems, towards polycentric ‘contemporary’ urban patterns, arranged following field models. Infrastructures have been traditionally on the spot for generations of architects and urban planners who, due to either their aesthetical condition, or an intuition of their organizational capacity, have been fascinated by the bursting of these elements in the territory. Projects which have explored the possibility to inhabit these huge structures, or to design with its language or scale for decades, have turned out to be always limited to morphological-typological issues. However, from the base of a research on ‘classic’ infrastructure –understood as the outcome of supplies and connectivity basic networks– a new nature of infrastructure is revealed, going beyond its known formal expression and exploring its potential to perform as a system of relationships, as a topological pattern, or as a flexible support for spontaneous and unanticipated activity. The breakthrough of a relational consciousness, as well as the modern concern about the generation of ‘a new city for a new society’ are shown, through the works of several authors and a selection of heterogeneous mostly-unbuilt projects. This semantic history of infrastructures, regarding not only typologies but also a broader concept able to operate in contemporary postmetropolitan territory, is drawn by following the paths of Fumihiko Maki in Investigations in Collective Form, Reyner Banham in Megastructure, or Shadrach Woods in The Man in the Street during the 60’s and 70’s and, lately, those of Stan Allen in Points + Lines, Edward Soja in Postmetropolis or Frei Otto in Occupying and Connecting. In order to do so, this thesis project sets, on one hand, a description of the urban context which can be assumed as contemporary, as well as its evolution and main features. On the other hand, complementary approaches help go deep into how this notion of infrastructural insight within the last decades has been created. An introduction describes the contextual conditions for this thesis project to become relevant: the relatively recent transit from a classic spatial occupation pattern, to the contemporary, disperse and polycentric pattern inherent to field models. A three-chapters core analyzes the infrastructures as the expression of an architecture of relations, as well as a support for a changing society and as a formless design logic. The last part, an epilogue, describes the urban contemporary territory and the suitability of the infrastructural logics to operate in it. A final conclusive section, lays the foundations for an open line of research on present functionality of these systems in architectural designs.

Spatial distribution of the soil carbon pool in a Holm oak dehesa in Spain

Aims Dehesas are agroforestry systems characterized by scattered trees among pastures, crops and/or fallows. A study at a Spanish dehesa has been carried out to estimate the spatial distribution of the soil organic carbon stock and to assess the influence of the tree cover. Methods The soil organic carbon stock was estimated from the five uppermost cm of themineral soil with high spatial resolution at two plots with different grazing intensities. The Universal Kriging technique was used to assess the spatial distribution of the soil organic carbon stocks, using tree coverage within a buffering area as an auxiliary variable. Results A significant positive correlation between tree presence and soil organic carbon stocks up to distances of around 8 m from the trees was found. The tree crown cover within a buffer up to a distance similar to the crown radius around the point absorbed 30 % of the variance in the model for both grazing intensities, but residual variance showed stronger spatial autocorrelation under regular grazing conditions. Conclusions Tree cover increases soil organic carbon stocks, and can be satisfactorily estimated by means of crown parameters. However, other factors are involved in the spatial pattern of the soil organic carbon distribution. Livestock plays an interactive role together with tree presence in soil organic carbon distribution.

Competition may explain the fine-scale spatial patterns and genetic structure of two co-occurring plant congeners

1. The spatial distribution of individual plants within a population and the population’s genetic structure are determined by several factors, like dispersal, reproduction mode or biotic interactions. The role of interspecific interactions in shaping the spatial genetic structure of plant populations remains largely unknown. 2. Species with a common evolutionary history are known to interact more closely with each other than unrelated species due to the greater number of traits they share. We hypothesize that plant interactions may shape the fine genetic structure of closely related congeners. 3. We used spatial statistics (georeferenced design) and molecular techniques (ISSR markers) to understand how two closely related congeners, Thymus vulgaris (widespread species) and T. loscosii (narrow endemic) interact at the local scale. Specific cover, number of individuals of both study species and several community attributes were measured in a 10 × 10 m plot. 4. Both species showed similar levels of genetic variation, but differed in their spatial genetic structure. Thymus vulgaris showed spatial aggregation but no spatial genetic structure, while T. loscosii showed spatial genetic structure (positive genetic autocorrelation) at short distances. The spatial pattern of T. vulgaris’ cover showed significant dissociation with that of T. loscosii. The same was true between the spatial patterns of the cover of T. vulgaris and the abundance of T. loscosii and between the abundance of each species. Most importantly, we found a correlation between the genetic structure of T. loscosii and the abundance of T. vulgaris: T. loscosii plants were genetically more similar when they were surrounded by a similar number of T. vulgaris plants. 5. Synthesis. Our results reveal spatially complex genetic structures of both congeners at small spatial scales. The negative association among the spatial patterns of the two species and the genetic structure found for T. loscosii in relation to the abundance of T. vulgaris indicate that competition between the two species may account for the presence of adapted ecotypes of T. loscosii to the abundance of a competing congeneric species. This suggests that the presence and abundance of close congeners can influence the genetic spatial structure of plant species at fine scales.

Use of DFOT Heat Pulse Method ForWetting Pattern Determination in Drip Irrigation Emitters

Although there are numerous accurate measuring methods to determine soil moisture content in a spot, until very recently there were no precise in situ and in real time methods that were able to measure soil moisture content along a line. By means of the Distributed Fiber Optic Temperature Measurement method or DFOT, the temperature in 0.12 m intervals and long distances (up to 10,000 m) with a high time frequency and an accuracy of +0.2º C is determined. The principle of temperature measurement along a fiber optic cable is based on the thermal sensitivity of the relative intensities of backscattered photons that arise from collisions with electrons in the core of the glass fiber. A laser pulse, generated by the DTS unit, traversing a fiber optic cable will result in backscatter at two frequencies. The DTS quantifies the intensity of these backscattered photons and elapsed time between the pulse and the observed returned light. The intensity of one of the frequencies is strongly dependent on the temperature at the point where the scattering process occurred. The computed temperature is attributed to the position along the cable from which the light was reflected, computed from the time of travel for the light.

Topological invariance and spatial scaling of surface roughness in two highly eroded zones of Mexico: a comparative study

The Fractal Image Informatics toolbox (Oleschko et al., 2008 a; Torres-Argüelles et al., 2010) was applied to extract, classify and model the topological structure and dynamics of surface roughness in two highly eroded catchments of Mexico. Both areas are affected by gully erosion (Sidorchuk, 2005) and characterized by avalanche-like matter transport. Five contrasting morphological patterns were distinguished across the slope of the bare eroded surface of Faeozem (Queretaro State) while only one (apparently independent on the slope) roughness pattern was documented for Andosol (Michoacan State). We called these patterns ?the roughness clusters? and compared them in terms of metrizability, continuity, compactness, topological connectedness (global and local) and invariance, separability, and degree of ramification (Weyl, 1937). All mentioned topological measurands were correlated with the variance, skewness and kurtosis of the gray-level distribution of digital images. The morphology0 spatial dynamics of roughness clusters was measured and mapped with high precision in terms of fractal descriptors. The Hurst exponent was especially suitable to distinguish between the structure of ?turtle shell? and ?ramification? patterns (sediment producing zone A of the slope); as well as ?honeycomb? (sediment transport zone B) and ?dinosaur steps? and ?corals? (sediment deposition zone C) roughness clusters. Some other structural attributes of studied patterns were also statistically different and correlated with the variance, skewness and kurtosis of gray distribution of multiscale digital images. The scale invariance of classified roughness patterns was documented inside the range of five image resolutions. We conjectured that the geometrization of erosion patterns in terms of roughness clustering might benefit the most semi-quantitative models developed for erosion and sediment yield assessments (de Vente and Poesen, 2005).

Spatial and temporal precipitation patterns of the Ebro River Basin, Spain

The Ebro River Basin, with around 85 000 km2 and located in NE Spain, is characterized by the high spatial heterogeneity of its geology, topography, climatology and land use. Rainfall is one of the most important climatic variables studied owing to its non-homogenous behaviour in event and intensity, which creates drought, water runoff and soil erosion with negative environmental and social consequences. In this work we characterized the rainfall variability pattern in the Ebro River Basin using universal multifractal (UM) analysis, which estimates the concentration of the data around the precipitation average (C1, codimension average), the degree of multiscaling behaviour in time (? index) and the maximum probable singularity in the rainfall distribution ( s). A spatial and temporal analysis of the UM parameters is applied to study the possible changes. With this porpoise, 60 daily rainfall series were selected from 132 synthetic series generated by Luna and Balairón (AEMet). These daily rainfall series present a length of 60 years, from 1950 to 2009. Each one of them was subdivided (1950?1970 and 1980?2009) to analyse the difference between the two periods. The range of variation of precipitation amounts and the frequency of dry events between both periods are discussed, as well as the evolution of the UM parameters through the years.

On the spatial distribution of woody plant species in a tropical montane cloud forest

Abstract The cloud forest is a special type of forest ecosystem that depends on suitable conditions of humidity and temperature to exist; hence, it is a very fragile ecosystem. The cloud forest is also one of the richest ecosystems in terms of species diversity and rate of endemism. However, today, it is one of the most threatened ecosystems in the world. Little is known about tree species distribution and coexistence among cloud forest trees. Trees are essential to understanding ecosystem functioning and maintenance because they support the ecosystem in important ways. For this dissertation, an analysis of woody plant species distribution at a small scale in a north-Peruvian Andean cloud forest was performed, and some of the factors implicated in the observed patterns were identified. Towards that end, different natural factors acting on species distribution within the forest were investigated: (i) intra-specific arrangements, (ii) heterospecific spatial relationships and (iii) relationships with external environmental factors. These analyses were conducted first on standing woody plants and then on seedlings. The woody plants were found to be clumped in the forest, either considering all the species together or each species separately. However, each species presented a specific pattern and specific spatial relationship among different-age individuals. Dispersal mode, growth form and shade tolerance played roles in the final distribution of the species. Furthermore, spatial associations among species, either positive or negative, were observed. These associations were more numerous when considering individuals of the interacting species at different developmental stages, i.e., younger individuals from one species and older individuals from another. Accordingly, competition and facilitation are asymmetric processes and vary throughout the life of an individual. Moreover, some species appear to prefer certain habitat conditions and avoid other habitats. The habitat definition that best explains species distribution is that which includes both environmental and stand characteristics; thus, a combination of these factors is necessary to understanding species' niche preferences. Seedling distribution was also associated with habitat conditions, but these conditions explained less than the 30% of the spatial variation. The position of conspecific adult individuals also affected seedling distribution; although the seedlings of many tree species avoid the vicinity of conspecifics, a few species appeared to prefer the formation of cohorts around their parent trees. The importance of habitat conditions and distance dependence with conspecifics varied among regions within the forest as well as on the developmental stage of the stand. The results from this thesis suggest that different species can coexist within a given space, forming a “puzzle” of species as a result of the intra- and interspecific spatial relationships along with niche preferences and adaptations that operate at different scales. These factors not only affect each species in a different way, but specific preferences also vary throughout species' lifespans. Resumen Resumen El bosque de niebla es uno de los ecosistemas más amenazados del mundo además de ser uno de los más frágiles. Son formaciones azonales que dependen de la existencia de unas condiciones de humedad y temperatura que permitan la formación de nubes que cubran el bosque; lo que dificulta en gran medida su conservación. También es uno de los ecosistemas con mayor riqueza de especies además de tener uno de los mayores porcentajes de endemismos. Uno de los aspectos más importantes para entender el ecosistema, es identificar y entender los elementos que lo componen y los mecanismos que regulan las relaciones entre ellos. Los árboles son el soporte del ecosistema. Sin embargo, apenas hay información sobre la distribución y coexistencia de los árboles en los bosques de niebla. Esta tesis presenta un análisis de la distribución a pequeña escala de las plantas leñosas en un bosque de niebla situado en la cordillera andina del norte de Perú; así como el análisis de algunos de los factores que pueden estar implicados en que se origine la distribución observada. Para este propósito se estudia cómo influyen factores de diferente naturaleza en la distribución de las especies (i) organización intra-específica (ii) relaciones espaciales heterospecíficas y (iii) relación con factores ambientales externos. En estos análisis se estudiaron primero las plantas jóvenes y las adultas, y después las plántulas. Los árboles aparecieron agregados en el bosque, tanto considerando todos a la vez como cuando se estudió cada especie por separado. Sin embargo, cada especie mostró un patrón distinto así como una particular relación espacial entre individuos jóvenes y adultos. El modo de dispersión, la forma de vida y la tolerancia de la especies estuvieron relacionados con el patrón general observado. Se vio también que ciertas especies aparecían relacionadas con otras, tanto de forma positiva (compartiendo zonas) como negativa (apareciendo en áreas distintas). Las asociaciones fueron mucho más numerosas cuando se consideraron los pares de especies en diferente estado de desarrollo, es decir, individuos jóvenes de una especie e individuos mayores de la otra. Eso indicaría que los procesos de competencia y facilitación son asimétricos y además varían durante la vida de la planta. Por otro lado, algunas especies aparecen preferentemente bajo ciertas condiciones de hábitat y evitan otras. La definición de hábitat a la que mejor responden las especies es cuando se incluyen tanto variables ambientales como de masa; así que ambos tipos de variables son necesarias para entender la preferencia de las especies por ciertos nichos. La distribución de las plántulas también estuvo relacionada con condiciones de hábitat, pero eso sólo llegaba a explicar hasta un 30% de la variabilidad espacial. La posición de los adultos de la misma especie también afectó a la distribución de las plántulas. En bastantes especies las plántulas evitan la cercanía de adultos de su misma especie, padres potenciales, aunque algunas especies aisladas mostraron el patrón contrario y aparecieron preferentemente en las mismas áreas que sus padres. La importancia de las condiciones de hábitat y posición de los adultos en la disposición de las plántulas varía de una zona a otra del bosque y además también varía según el estado de desarrollo de la masa.