Elementos peatonales de las ciudades medias españolas : tipos, orígenes, relaciones y articulaciones

A través de esta tesis se busca aportar un análisis de los elementos peatonales que cubren las necesidades y las infraestructuras de apoyo al peatón, siendo un campo poco desarrollado en nuestro país a diferencia de otros como Dinamarca, Reino Unido, Alemania y Holanda, donde las actuaciones peatonales ejecutadas son tradicionales y muy exitosas. Se trata por tanto de un tema poco estudiado pero que adquiere singular importancia en estos años de búsqueda de nuevos modelos urbanos que eviten el derroche de recursos, y avancen hacia formas de movilidad más adecuadas a un mundo limitado y vulnerable. La investigación tiene un carácter eminentemente descriptivo cuyo principal valor es documentar y analizar sistemáticamente una realidad poco conocida, aportándose datos para entender el origen y desarrollo de las actuaciones peatonales. El propósito es ayudar a cimentar este campo en nuestro país, definiendo con precisión el estado de la cuestión geográficamente y dejando para un futuro una cartografía temática completa. El objeto general es estudiar y comprender los elementos peatonales de un número limitado de ciudades medias españolas, marcándose como objetivos específicos los siguientes: - Identificar y describir los principales elementos que conforman la red peatonal de las ciudades. ‐ Estudiar las tipologías espaciales dominantes, sus características y orígenes, pudiendo distinguir de un lado, las desarrolladas en la ciudad consolidada, como las actuaciones en los cascos antiguos; y por otro, las ejecutadas en la ciudad nueva a través de fórmulas arquitectónicas generalmente residenciales con espacios peatonales adjuntos. ‐ Analizar los referidos elementos para comprobar si han conformado redes de uso peatonal, si se han constituido articulando la ciudad como en el caso de los ejes lineales en las ciudades con río o mar, o si han influenciado el carácter del entorno en el que se han realizado, centrando el interés en los cascos históricos peatonalizados. ‐ Comprobar si hay actuaciones complementarias a los procesos de urbanización y mejora urbanística, así como la aprobación de ayudas y subvenciones para afrontar las mismas. ‐ Finalmente, describir y evaluar los procesos y procedimientos mediante los que se produce la peatonalización de determinadas áreas urbanas, los orígenes, los instrumentos y fechas de su puesta en marcha, aportando una cronología peatonal. La metodología comienza con la acotación de 22 ciudades españolas, capitales de provincia con una población entre 100.000 y 500.000 habitantes según los datos del INE del año 2009. La distribución es equilibrada con la selección de ciudades de costa, de interior, con río/ría, con carácter patrimonial, ciudades en islas, etc. El análisis de los elementos peatonales de estas urbes se realiza gracias a la gran resolución de las bases cartográficas públicas y privadas que hay en la actualidad en internet, especialmente con el uso de las herramientas de Google Earth y Google Maps en su versión “street view”, softwares que permiten un paseo virtual de cada ciudad detallándose con gran precisión las actuaciones peatonales. Las ciudades elegidas son: ‐ 11 ciudades costeras: Alicante, Almería, Bilbao, Cádiz, Castellón de la Plana, Huelva, Las Palmas de Gran Canaria, Palma de Mallorca, Santander, San Sebastián y Tarragona. ‐ 11 ciudades de interior: Albacete, Burgos, Córdoba, León, Lérida, Logroño, Oviedo, Murcia, Pamplona, Valladolid y Vitoria. Finalmente, por tener los porcentajes mas elevados de las áreas peatonales de sus casco histórico, se eligen 3 casos de ciudades representativas para un desarrollo peatonal en profundidad, seleccionándose Burgos, San Sebastián y Pamplona. Otros aspectos metodológicos interesantes a destacar, es la extensa documentación adquirida a través de las fuentes de Internet, sobre todo a la hora de desarrollar el último capítulo “Procesos de peatonalización de las ciudades españolas” centrado en las 3 ciudades elegidas. Se consulta las hemerotecas de periódicos locales, los boletines oficiales, las actas de los plenos de los Ayuntamientos y los documentos de planificación (Planes Generales, Planes Especiales, Planes y Proyectos Municipales, Planes de Movilidad), adquiriéndose una precisa información en cuanto a la cronología de las fechas y extensión del trabajo. Se complementa la metodología con las entrevistas a actores claves en el proceso peatonal, como el personal técnico de los Ayuntamiento y los técnicos redactores de los proyectos; y con la colaboración directa de los técnicos de los consistorios de las 3 ciudades seleccionadas, aportando todo tipo de información demandada. En la tesis, se introduce los elementos peatonales mediante el estudio de los espacios peatonales y los procesos de peatonalización existentes, a través del análisis de la literatura consultada aportándose información valiosa en cuanto a la definición de los modelos peatonales, exponiéndose una breve historia de las peatonalizaciones. Destacan las publicaciones de los expertos europeos en la materia como los autores Carmen Hass-Klau, Rolf Monheim, Rob Krier y Collin Buchanan. En España, las actuaciones peatonales fueron tardías en el tiempo en comparación con el resto de Europa, con casi 40 años de retraso. Las referencias bibliográficas españolas en esta materia son escasas siendo las mas significativas las realizadas por Alfonso Sanz y por el profesor Julio Pozueta destacando una de sus últimas publicaciones “La ciudad paseable” describiéndose los elementos peatonales existentes en las ciudades, sirviendo como modelo y ejemplo para esta investigación. En base a lo anterior, se definen los elementos peatonales como aquellos espacios públicos libres de edificación que, por sus características y diseño, garantizan un confortable uso estancial, de tránsito peatonal y están expresamente reservados para ello. En la tesis se detectan y se especifican los siguientes elementos peatonales: las calles y plazas peatonales, las aceras y bulevares de una anchura superior a 8 metros, los paseos peatonales, los elementos de paso (tradicionales y mecánicos), espacios peatonales ligados a nuevas áreas residenciales, los soportales y pasajes peatonales con una anchura mínima de 4 metros, recintos feriales, espacios libres y zonas verdes y parques. Para cada una de las 22 ciudades seleccionadas se realiza un exhaustivo inventario de los anteriores elementos a través de fichas, aportando un gran número de proyectos individuales ejecutados en cada urbe. Esta información se recoge gráficamente en 2 fotoplanos, conformándose como herramientas fundamentales para esta investigación. El Fotoplano 1 se hace sobre la base de la ciudad del Google Earth diferenciándose las áreas peatonales y las zonas verdes. En el Fotoplano 2, y con el soporte anterior, se distinguen las tipologías persistentes y dominantes como son los grandes ejes peatonales, las redes peatonales locales y los elementos peatonales singulares. Finalmente, a partir de los 2 fotoplanos, se realiza la denominada “Huella peatonal” que destaca por su atractivo y pregnancia, algo de lo que el campo de lo peatonal es muy útil. A través de la huella se obtiene una rápida información al tener el plano dos colores, color naranja para lo peatonal (formado por las áreas peatonales y zonas verdes) y color negro para el resto el resto no peatonal. Toda esta documentación gráfica y descriptiva de los elementos peatonales de cada ciudad, queda ampliamente recogida en el Anexo de la tesis. La “Huella peatonal” se considera como el plano básico en la investigación obteniendo una percepción rápida, clara y visual del espacio peatonal de cada ciudad. Se arroja información morfológica con respecto a la distribución espacial, articulación, equilibrio, modulación, cohesión, concentración peatonal, etc. de cada una de las ciudades seleccionadas. A través de la huella se resuelve muchos de las cuestiones enmarcadas como objetivos de la tesis, comprobándose de forma general, que los elementos peatonales dentro de la trama urbana están desarticulados ya que no hay una estructura que module la ciudad como un conjunto cohesionado. Se localizan pequeñas redes conectadas entre sí, que de forma heterogénea, están dispersas por la ciudad. Se verifica que los elementos peatonales con mayor concentración y presencia en las ciudades analizadas son los ejes peatonales, los cascos históricos y los elementos peatonales en las nuevas áreas residenciales. En la investigación, además de estos 3 elementos mencionados, se analiza con mayor profundidad otros tipos que destacan por su persistencia y repetición en la “Huella peatonal” de cada una de las 22 ciudades, como son las plazas peatonales, los soportales y los elementos singulares peatonales: espacios tradicionales (recintos feriales, pasajes, puentes históricos), nuevos elementos peatonales (pasarelas peatonales, elementos mecánicos de paso), áreas peatonales entorno a edificios culturales y otros casos como los consistentes en la recuperación y transformación de edificaciones y construcciones urbanas de diferente uso y tipología, en nuevas áreas peatonales. Se aporta luz sobre su definición, localización, cronología, proporción, presencia y génesis, facilitando resultados de cada uno de ellos. De forma paralela, se obtiene información de índole cuantitativa a través de las mediciones de la huella, obteniéndose datos y porcentajes de extensión de la ciudad, de las zonas peatonales y de las zonas verdes, de los cascos históricos y de sus superficies peatonales, de los metros lineales de dichos cascos históricos y de sus calles peatonales. Se proporcionan ratios e índices que se consideran fundamentales para una mejor comprensión peatonal de cada urbe. Se establece una clasificación de las ciudades en cada uno de los ámbitos descritos, destacando las ciudades de Pamplona, San Sebastián y Burgos por arrojar las cifras peatonales más positivas encabezando la mayoría de las tablas clasificatorias. La peor ciudad valorada es Alicante, seguida según criterio, de Almería, Palma de Mallorca y Las Palmas de GC. En el capítulo final de la investigación, se eligen las ciudades de Burgos, San Sebastián y Pamplona como casos de estudio representativos. El objetivo es obtener un conocimiento más preciso en cuanto a su origen, cronología y procesos de las peatonalizaciones. Cada ciudad se compone de un cuadro cronológicos por etapas, desarrollándose de forma exhaustiva cada una de ellas, enunciadas normalmente, como instrumentos de planeamiento o proyectos de urbanización, acompañadas de los planos de las actuaciones peatonales. Al final de cada etapa se aportan datos de las cifras de las peatonalizaciones que se van realizando, con un cuadro numérico de localizaciones, superficie peatonal, superficie reurbanizada, longitud peatonal y el grado de peatonalización del casco histórico, además de los datos acumulados por etapa. De las conclusiones cruzadas de estas 3 ciudades, destaca la comprobación de la tendencia de ir peatonalizando los centros antiguos llegando incluso al 100% de calles peatonales en el caso de San Sebastián. No obstante, es difícil confirmar si el objetivo es una peatonalización global de los cascos, ya que son muchas las variables que afectan e influyen en el proceso peatonal de cada ciudad. Se coteja adicionalmente cómo se extiende la cultura peatonal desde el interior al exterior, mas allá de los cascos históricos hacia los ensanches próximos. En cuanto al origen de las peatonalizaciones, se concluye que no hay una concepción inicial y teórica marcada, a partir de la cual se va desarrollando la trama peatonal de cada ciudad. Se puede afirmar que los procesos peatonales ejecutados son variados y de diversa índole, sin poder precisar un hecho como causa principal. Según el momento y las circunstancias, las peatonalizaciones responden a distintas iniciativas promovidas por los comerciantes, por las administraciones locales o por los técnicos de planeamiento, con sus expertas aportaciones en los planes. Estos tres agentes actúan como impulsores de proyectos individuales peatonales que se van solapando en el tiempo. Como punto final de la tesis, se propone una serie de nuevas líneas de investigación que pueden servir como estudio adicional y complementario, respondiendo a aspectos tan relevantes como cuestiones de índole económica, social y de movilidad, fuertemente ligadas a las peatonalizaciones, tal y como se demuestra en la literatura consultada. ABSTRACT This thesis aims to analyse the pedestrian elements dealing with the needs and infrastructures that support pedestrians, as this is a field that is little developed in our country, in comparison to other regions such as Denmark, the United Kingdom, Germany and the Netherlands, where the pedestrian actions implemented are traditional and highly successful. Therefore, even though little research has been done about this topic, this field is increasingly important throughout these years of search for urban models intended to prevent the waste of resources and to develop new mobility ways, more adapted to a limited, vulnerable world. This research is essentially descriptive and it mainly aims to document and systematically analysed a scarcely known reality by providing data in order to understand the origin and development of pedestrian actions. This thesis intends to lay the foundations of this field in Spain, accurately defining the state of the art from a geographical point of view and preparing a full thematic map that may be used in the future. The overall aim is to study and understand the pedestrian elements of a limited number of Spanish medium-size cities, establishing the following specific goals: - To identify and describe the main elements comprising cities' pedestrian networks. - To study of special predominant typologies, their features and origins, with the possibility to, on the one hand, make a difference between the typologies developed in well-established cities, such as the actions in historic quarters, and, on the other hand, those implemented in new cities through generally residential architectural formulae with adjoining pedestrian areas. - To analyse the said elements in order to check whether they have resulted in pedestrian-use networks, whether they have been established by organising the city, such as the linear axes in coastal and riverside cities, or whether they have had an impact on the character of the areas where these elements have been implemented, focusing on pedestrian historic quarters. - Check whether there are actions supplementary to the urban development and urban improvement processes, as well as the approval of financial support and subventions to deal with these actions. - Finally, to describe and assess the processes and procedures by which the pedestrianisation of certain urban areas is carried out, the origins, the instruments and the date of their implementation, providing a pedestrian timeline. The methodology starts by defining 22 Spanish cities, province capitals with a population ranging from 100,000 to 500,000 inhabitants, according to the data recorded by the Spanish Statistics Institute (INE) in 2009. The distribution of coastal, riverside and interior cities, as well as of patrimonial cities and cities in islands, etc. is well balanced. The analysis of the pedestrian elements of these cities is made with the great resolution of the public and private map databases that can be accessed on the Internet, especially using the "street-view" version of tools such as Google Earth and Google Maps, software applications that allow to go for a virtual walk in each city, providing highly precise details about the pedestrian actions. The following cities have been chosen: ‐ 11 coastal cities: Alicante, Almería, Bilbao, Cádiz, Castellón de la Plana, Huelva, Las Palmas de Gran Canaria, Palma de Mallorca, Santander, San Sebastián and Tarragona. ‐ 11 interior cities: Albacete, Burgos, Córdoba, León, Lérida, Logroño, Oviedo, Murcia, Pamplona, Valladolid and Vitoria. Finally, as they have the highest percentages regarding the pedestrian areas found in the historic quarters, 3 cities representing a deep pedestrian development have been chosen: Burgos, San Sebastián and Pamplona. Other significant methodological aspects are the many documents found from online sources, especially when preparing the last chapter: “Processes for the pedestrianisation of Spanish cities”, which focuses on the 3 cities chosen. Local newspaper and periodical libraries, official gazettes, the minutes of plenary sessions of councils and the zoning regulation documents (General Zoning Plans, Special Zoning Plans, Municipal Plans and Projects, Mobility Plans, etc.) have been consulted, obtaining accurate information regarding the timeline and the extension of the works carried out. The methodology is supplemented by interviews with key players in the pedestrianisation process, such as the technical staff in councils and the officers drafting and designing the projects, as well as with the direct collaboration by the officers of the councils of the 3 cities chosen, who provided all the information requested. The thesis introduces the pedestrian elements by studying pedestrian areas and existing pedestrianisation processes through the analysis of the literature consulted, providing valuable information for the definition of pedestrian models and showing a brief history and background of the pedestrianisation process. Remarkable papers published by some European experts in the field, such as Carmen Hass-Klau, Rolf Monheim and Collin Buchanan, are covered in the thesis. In Spain, the pedestrianisation actions were late in time in comparison to the rest of Europe, with at least 40 years of delay. The Spanish literature references in this field are limited. The most significant papers are those published by Alfonso Sanz and by Professor Julio Pozueta, with a special mention of one of his last works "La ciudad paseable" (The Walkable City), which describes the pedestrian elements found in the cities and is used a model and an example for this research. Based on the elements above, pedestrian elements are defined as those construction-free public areas that, due to their features and design, ensure the comfortable, convenient use of the spaces, characterised by pedestrian traffic and specifically reserved for this purpose. The thesis detects and specifies the following pedestrian elements: pedestrian streets and squares, pavements (sidewalks) and boulevards with a width exceeding 8 metres, pedestrian promenades, crossing elements (traditional and mechanical), pedestrian spaces linked to new residential areas, colonnades and pedestrian passages or narrow streets with a minimum width of 4 metres, exhibition sites, free spaces and green areas and parks. For each of the 22 cities chosen, a thorough inventory of the elements mentioned above has been made by using worksheets, providing a significant number of individual projects developed in each city. This information is graphically collected and displayed on 2 photomaps, resulting in tools essential for this research. Photomap 1 is made based on the city displayed by Google Earth, making a difference between pedestrian areas and green areas. On Photomap 2, using the tool mentioned above, a difference can be made between persistent and predominant typologies, such as the big pedestrian axes, the local pedestrian networks and singular pedestrian elements. Finally, the 2 photomaps are used in order to establish the so-called "pedestrian footprint", which is highlighted by its attractiveness and appeal, concepts for which the pedestrian field is very useful. With the pedestrian footprint, quick information can be obtained, since the map shows two colours: orange for pedestrian elements (made up of pedestrian areas and green areas) and black for the other non-pedestrian elements. A significant part of these graphic, descriptive documents about each city's pedestrian elements can be found in the thesis appendix. The "Pedestrian Footprint" is regarded in the research as the basic map, obtaining a quick, clear and visual perception of each city's pedestrian space. This footprint provides morphological information regarding the space distribution, the organisation, the balance, the modulation, the cohesion, the pedestrian concentration, etc. in each of the cities chosen. The pedestrian footprint helps solve many of the questions established as the thesis goals, proving that, in general, the pedestrian elements are not organised in the urban plot, as there is no structure modulating the city as a properly linked set of elements. Small networks linked to each other and heterogeneously scattered all over the city are found. It has been verified that the pedestrian elements with the highest concentration and presence in the cities analysed are the pedestrian axes, the historic quarters and the pedestrian elements found in the new residential areas. Besides these 3 elements mentioned, the research analyses in depth other types that are remarkable due to their persistence and repetition in the "Pedestrian Footprint" of each of the 22 cities, such as the pedestrian squares, the colonnades and the singular pedestrian elements: traditional spaces (exhibition sites, passages, historic bridges), new pedestrian elements (pedestrian footbridges, mechanical crossing elements), pedestrian areas around cultural buildings and other cases such as those consisting of recovering and transforming building and urban constructions, intended for a wide range of purposes and of different types, into new pedestrian areas. This work puts light on the definition, location, timeline, proportion, presence and origin, providing results for each of these concepts. At the same time, quantitative information is obtained by measuring the footprint, getting data and percentages on the size of the city, the pedestrian areas and the green areas, the historic quarters and the pedestrian zones, the linear metres of such historic quarters and pedestrian streets. The footprint also provides ratios and rates that are considered as essential in order to better understand the pedestrian elements of each city. A classification of cities is established for each of the areas described, highlighting the cities of Pamplona, San Sebastián and Burgos, as they provide the most positive pedestrian figures and lead most of the classification tables or rankings. According to the criteria considered, the city with the worst values is Alicante, followed by Almería, Palma de Mallorca and Las Palmas de Gran Canaria. In the final chapter in the thesis, the cities of Burgos, San Sebastián and Pamplona are chosen as representative study cases. The aim is to gain more accurate knowledge regarding the pedestrianisation origin, timeline and processes. Each city comprises a chronological sequence made of stages, each of which is thoroughly developed and usually announced as zoning plans or urban development projects, accompanied by the plans of the pedestrian actions. At the end of each stage, data with the figures of the pedestrianisation projects are provided, including a numerical chart with locations, pedestrian area, redeveloped urban areas, pedestrian length and pedestrianisation degree in the historic quarter, as well as the data cumulated throughout each stage. In the crossed conclusions on the three cities, the trend of gradually pedestrianising the historic quarters (even reaching the 100% of the pedestrian streets as in San Sebastián) is verified. However, it is difficult to confirm whether the purpose is to reach an overall pedestrianisation of the historic quarters, since there are many variables that affect and influence each city's pedestrianisation process. In addition, the spread of the pedestrian culture from the internal areas to the external areas, beyond the historic quarters to the nearby expansion areas, is compared. Regarding the origin of pedestrianisations, the thesis comes to the conclusion that there is no initial or theoretical conception from which each city's pedestrian plot is developed. The pedestrian processes implemented are varied and diverse but no fact can be specified as the main cause. Based on the moment and circumstances, the pedestrianisation processes are due to different initiatives promoted by shopkeepers, by the local administrations or by the zoning officers, with their expert contributions to the plans. These three players promote and drive individual pedestrianisation projects that overlap in the course of time. Finally, new lines of research are put forwards, as they can be taken as an additional study and as a supplement to this research, dealing with aspects as significant as the economic, social and mobility factors, closely linked to the pedestrianisation processes, as proven in the literature consulted.

Residual delay maps unveil global patterns of atmospheric nonlinearity and produce improved local forecasts

We use residual-delay maps of observational field data for barometric pressure to demonstrate the structure of latitudinal gradients in nonlinearity in the atmosphere. Nonlinearity is weak and largely lacking in tropical and subtropical sites and increases rapidly into the temperate regions where the time series also appear to be much noisier. The degree of nonlinearity closely follows the meridional variation of midlatitude storm track frequency. We extract the specific functional form of this nonlinearity, a V shape in the lagged residuals that appears to be a basic feature of midlatitude synoptic weather systems associated with frontal passages. We present evidence that this form arises from the relative time scales of high-pressure versus low-pressure events. Finally, we show that this nonlinear feature is weaker in a well regarded numerical forecast model (European Centre for Medium-Range Forecasts) because small-scale temporal and spatial variation is smoothed out in the grided inputs. This is significant, in that it allows us to demonstrate how application of statistical corrections based on the residual-delay map may provide marked increases in local forecast accuracy, especially for severe weather systems.

Multiple-complete-digest restriction fragment mapping: Generating sequence-ready maps for large-scale DNA sequencing

Multiple-complete-digest mapping is a DNA mapping technique based on complete-restriction-digest fingerprints of a set of clones that provides highly redundant coverage of the mapping target. The maps assembled from these fingerprints order both the clones and the restriction fragments. Maps are coordinated across three enzymes in the examples presented. Starting with yeast artificial chromosome contigs from the 7q31.3 and 7p14 regions of the human genome, we have produced cosmid-based maps spanning more than one million base pairs. Each yeast artificial chromosome is first subcloned into cosmids at a redundancy of ×15–30. Complete-digest fragments are electrophoresed on agarose gels, poststained, and imaged on a fluorescent scanner. Aberrant clones that are not representative of the underlying genome are rejected in the map construction process. Almost every restriction fragment is ordered, allowing selection of minimal tiling paths with clone-to-clone overlaps of only a few thousand base pairs. These maps demonstrate the practicality of applying the experimental and software-based steps in multiple-complete-digest mapping to a target of significant size and complexity. We present evidence that the maps are sufficiently accurate to validate both the clones selected for sequencing and the sequence assemblies obtained once these clones have been sequenced by a “shotgun” method.

Congruent Docking of Dimeric Kinesin and ncd into Three-dimensional Electron Cryomicroscopy Maps of Microtubule–Motor ADP Complexes

We present a new map showing dimeric kinesin bound to microtubules in the presence of ADP that was obtained by electron cryomicroscopy and image reconstruction. The directly bound monomer (first head) shows a different conformation from one in the more tightly bound empty state. This change in the first head is amplified as a movement of the second (tethered) head, which tilts upward. The atomic coordinates of kinesin·ADP dock into our map so that the tethered head associates with the bound head as in the kinesin dimer structure seen by x-ray crystallography. The new docking orientation avoids problems associated with previous predictions; it puts residues implicated by proteolysis-protection and mutagenesis studies near the microtubule but does not lead to steric interference between the coiled-coil tail and the microtubule surface. The observed conformational changes in the tightly bound states would probably bring some important residues closer to tubulin. As expected from the homology with kinesin, the atomic coordinates of nonclaret disjunctional protein (ncd)·ADP dock in the same orientation into the attached head in a map of microtubules decorated with dimeric ncd·ADP. Our results support the idea that the observed direct interaction between the two heads is important at some stages of the mechanism by which kinesin moves processively along microtubules.

Coexistence of linear zones and pinwheels within orientation maps in cat visual cortex

Revealing the layout of cortical maps is important both for understanding the processes involved in their development and for uncovering the mechanisms underlying neural computation. The typical organization of orientation maps in the cat visual cortex is radial; complete orientation cycles are mapped around orientation singularities. In contrast, long linear zones of orientation representation have been detected in the primary visual cortex of the tree shrew. In this study, we searched for the existence of long linear sequences and wide linear zones within orientation preference maps of the cat visual cortex. Optical imaging based on intrinsic signals was used. Long linear sequences and wide linear zones of preferred orientation were occasionally detected along the border between areas 17 and 18, as well as within area 18. Adjacent zones of distinct radial and linear organizations were observed across area 18 of a single hemisphere. However, radial and linear organizations were not necessarily segregated; long (7.5 mm) linear sequences of preferred orientation were found embedded within a typical pinwheel-like organization of orientation. We conclude that, although the radial organization is dominant, perfectly linear organization may develop and perform the processing related to orientation in the cat visual cortex.

A novel modifier gene for plasma von Willebrand factor level maps to distal mouse chromosome 11

Type 1 von Willebrand disease (VWD), characterized by reduced levels of plasma von Willebrand factor (VWF), is the most common inherited bleeding disorder in humans. Penetrance of VWD is incomplete, and expression of the bleeding phenotype is highly variable. In addition, plasma VWF levels vary widely among normal individuals. To identify genes that influence VWF level, we analyzed a genetic cross between RIIIS/J and CASA/Rk, two strains of mice that exhibit a 20-fold difference in plasma VWF level. DNA samples from F2 progeny demonstrating either extremely high or extremely low plasma VWF levels were pooled and genotyped for 41 markers spanning the autosomal genome. A novel locus accounting for 63% of the total variance in VWF level was mapped to distal mouse chromosome 11, which is distinct from the murine Vwf locus on chromosome 6. We designated this locus Mvwf for “modifier of VWF.” Additional genotyping of as many as 2407 meioses established a high resolution genetic map with gene order Cola1-Itg3a-Ngfr-Mvwf/Gip-Hoxb9-Hoxb1-Cbx·rs2-Cox5a-Gfap. The Mvwf candidate interval between Ngfr and Hoxb9 is ≈0.5 centimorgan (cM). These results demonstrate that a single dominant gene accounts for the low VWF phenotype of RIIIS/J mice in crosses with several other strains. The pattern of inheritance suggests a gain-of-function mutation in a unique component of VWF biosynthesis or processing. Characterization of the human homologue for Mvwf may have relevance for a subset of type 1 VWD cases and may define an important genetic factor modifying penetrance and expression of mutations at the VWF locus.

Formation of temporal-feature maps by axonal propagation of synaptic learning

Computational maps are of central importance to a neuronal representation of the outside world. In a map, neighboring neurons respond to similar sensory features. A well studied example is the computational map of interaural time differences (ITDs), which is essential to sound localization in a variety of species and allows resolution of ITDs of the order of 10 μs. Nevertheless, it is unclear how such an orderly representation of temporal features arises. We address this problem by modeling the ontogenetic development of an ITD map in the laminar nucleus of the barn owl. We show how the owl's ITD map can emerge from a combined action of homosynaptic spike-based Hebbian learning and its propagation along the presynaptic axon. In spike-based Hebbian learning, synaptic strengths are modified according to the timing of pre- and postsynaptic action potentials. In unspecific axonal learning, a synapse's modification gives rise to a factor that propagates along the presynaptic axon and affects the properties of synapses at neighboring neurons. Our results indicate that both Hebbian learning and its presynaptic propagation are necessary for map formation in the laminar nucleus, but the latter can be orders of magnitude weaker than the former. We argue that the algorithm is important for the formation of computational maps, when, in particular, time plays a key role.

Imaging neuroscience: Principles or maps?

This article reviews some recent trends in imaging neuroscience. A distinction is made between making maps of functional responses in the brain and discerning the rules or principles that underlie their organization. After considering developments in the characterization of brain imaging data, several examples are presented that highlight the context-sensitive nature of neuronal responses that we measure. These contexts can be endogenous and physiological, reflecting the fact that each cortical area, or neuronal population, expresses its dynamics in the context of interactions with other areas. Conversely, these contexts can be experimental or psychological and can have a profound effect on the regional effects elicited. In this review we consider experimental designs and analytic strategies that go beyond cognitive subtraction and speculate on how functional imaging can be used to address both the details and principles underlying functional integration and specialization in the brain.

Gene for the catalytic subunit of mouse DNA-dependent protein kinase maps to the scid locus.

The gene encoding the catalytic subunit of DNA-dependent protein kinase (DNA-PKcs) has been proposed recently as a candidate gene for the mouse severe combined immune deficiency (scid) locus. We have used a partial cDNA clone for human DNA-PKcs to map the mouse homologue using a large interspecific backcross panel. We found that the mouse gene for DNA-PKcs does not recombine with scid, consistent with the hypothesis that scid is a mutation in the mouse gene for DNA-PKcs.

Gene for the catalytic subunit of the human DNA-activated protein kinase maps to the site of the XRCC7 gene on chromosome 8.

The DNA-activated serine/threonine protein kinase (DNA-PK) is composed of a large (approximately 460 kDa) catalytic polypeptide (DNA-PKcs) and Ku, a heterodimeric DNA-binding component (p70/p80) that targets DNA-PKcs to DNA. A 41-kbp segment of the DNA-PKcs gene was isolated, and a 7902-bp segment was sequenced. The sequence contains a polymorphic Pvu II restriction enzyme site, and comparing the sequence with that of the cDNA revealed the positions of nine exons. The DNA-PKcs gene was mapped to band q11 of chromosome 8 by in situ hybridization. This location is coincident with that of XRCC7, the gene that complements the DNA double-strand break repair and V(D)J recombination defects (where V is variable, D is diversity, and J is joining) of hamster V3 and murine severe combined immunodeficient (scid) cells.