Efecto de las cubiertas ajardinadas sobre el microclima urbano de verano

El cambio climático y sus efectos requieren con urgencia el desarrollo de estrategias capaces no solo de mitigar pero también permitir la adaptación de los sistemas afectados por este fenómeno a los cambios que están provocando a nivel mundial. Olas de calor más largas y frecuentes, inundaciones, y graves sequías aumentan la vulnerabilidad de la población, especialmente en asentamientos urbanos. Este fenómeno y sus soluciones potenciales han sido ampliamente estudiados en las últimas décadas desde diferentes perspectivas y escalas que analizan desde el fenómeno regional de isla de calor al aumento de la intensidad energética necesaria en los edificios para mantener las condiciones de confort en los escenarios de calentamiento que se predicen. Su comprensión requiere el entendimiento de este fenómeno y un profundo análisis de las estrategias que pueden corregirlo y adaptarse a él. En la búsqueda de soluciones a este problema, las estrategias que incorporan sistemas naturales tales como las cubiertas ajardinadas, las fachadas vegetadas y bosques urbanos, se presentan como opciones de diseño capaces de proporcionan múltiples servicios al ecosistema urbano y de regular y hacer frente a los efectos del cambio climático. Entre los servicios que aportan estos sistemas naturales se incluyen la gestión de agua de tormentas, el control del efecto isla de calor, la mejora de la calidad del aire y del agua, el aumento de la diversidad, y como consecuencia de todo lo anterior, la reducción de la huella ecológica de las ciudades. En la última década, se han desarrollado múltiples estudios para evaluar y cuantificar los servicios al ecosistema proporcionados por las infraestructuras verdes, y específicamente las cubiertas ajardinadas, sin embargo, determinados servicios como la capacidad de la regulación del microclima urbano no ha sido apenas estudiados. La mayor parte de la literatura en este campo la componen estudios relacionados con la capacidad de las cubiertas ajardinadas de reducir el efecto de la isla de calor, en una escala local, o acerca de la reducción de la demanda energética de refrigeración debida a la instalación de cubiertas ajardinadas en la escala de edificio. La escala intermedia entre estos dos ámbitos, la calle, desde su ámbito habitable cercano al suelo hasta el límite superior del cañón urbano que configura, no han sido objeto detallado de estudio por lo que es esta escala el objeto de esta tesis doctoral. Esta investigación tiene como objeto contribuir en este campo y aportar un mayor entendimiento a través de la cuantificación del impacto de las cubiertas ajardinadas sobre la temperatura y humedad en el cañón urbano en la escala de calle y con un especial foco en el nivel peatonal. El primer paso de esta investigación ha sido la definición del objeto de estudio a través del análisis y revisión de trabajos tanto teóricos como empíricos que investigan los efectos de cubiertas ajardinadas en el entorno construido, entendidas como una herramienta para la adaptación y mitigación del impacto del cambio climático en las ciudades. La literatura analizada, revela el gran potencial de los sistemas vegetales como herramientas para el diseño pasivo puesto que no solo son capaces de mejorar las condiciones climáticas y microclimaticas en las ciudades reduciendo su demanda energética, sino también la necesidad de mayor análisis en la escala de calle donde confluyen el clima, las superficies urbanas y materiales y vegetación. Este análisis requiere una metodología donde se integren la respuesta térmica de edificios, las variaciones en los patrones de viento y radiación, y la interacción con la vegetación, por lo que un análisis cuantitativo puede ayudar a definir las estrategias más efectivas para lograr espacios urbanos más habitables. En este contexto, el objetivo principal de esta investigación ha sido la evaluación cuantitativa del impacto de la cubierta ajardinada en el microclima urbano a escala de barrio en condiciones de verano en los climas mediterráneos continentales. Para el logro de este objetivo, se ha seguido un proceso que persigue identificar los modelos y herramientas de cálculo capaces de capturar el efecto de la cubierta ajardinada sobre el microclima, identificar los parámetros que potencian o limitan este efecto, y cuantificar las variaciones que microclima creado en el cañón urbano produce en el consumo de energía de los edificios que rodean éste espacio. La hipótesis principal detrás de esta investigación y donde los objetivos anteriores se basan es el siguiente: "una cubierta ajardinada instalada en edificios de mediana altura favorece el establecimiento de microclimas a nivel peatonal y reduce las temperaturas en el entorno urbano donde se encuentra”. Con el fin de verificar la hipótesis anterior y alcanzar los objetivos propuestos se ha seguido la siguiente metodología: • definición del alcance y limitaciones del análisis • Selección de las herramientas y modelos de análisis • análisis teórico de los parámetros que afectan el efecto de las cubiertas ajardinadas • análisis experimental; • modelización energética • conclusiones y futuras líneas de trabajo Dada la complejidad de los fenómenos que intervienen en la generación de unas determinadas condiciones microclimáticas, se ha limitado el objeto de este estudio a las variables de temperatura y humedad, y sólo se han tenido en cuenta los componentes bióticos y abióticos del sistema, que incluyen la morfología, características superficiales del entorno estudiado, así como los elementos vegetales. Los componentes antrópicos no se han incluido en este análisis. La búsqueda de herramientas adecuadas para cumplir con los objetivos de este análisis ha concluido en la selección de ENVI-met v4 como el software más adecuado para esta investigación por su capacidad para representar los complejos fenómenos que caracterizan el microclima en cañones urbanos, en una escala temporal diaria y con unas escala local de vecindario. Esta herramienta supera el desafío que plantean los requisitos informáticos de un cálculo completo basado en elementos finitos realizados a través de herramientas de dinámica de fluidos computacional (CFD) que requieren una capacidad de cálculo computacional y tiempo privativos y en una escala dimensional y temporal limitada a esta capacidad computacional lo que no responde a los objetivos de esta investigación. ENVI-met 4 se basa es un modelo tridimensional del micro clima diseñado para simular las interacciones superficie-planta-aire en entornos urbanos. Basado en las ecuaciones fundamentales del equilibrio que representan, la conservación de masa, energía y momento. ENVI-met es un software predictivo, y como primer paso ha requerido la definición de las condiciones iniciales de contorno que se utilizan como punto de partida por el software para generar su propio perfil de temperatura y humedad diaria basada en la localización de la construcción, geometría, vegetación y las superficies de características físicas del entorno. La geometría de base utilizada para este primer análisis se ha basado en una estructura típica en cuanto al trazado urbano situada en Madrid que se ha simulado con una cubierta tradicional y una cubierta ajardinada en sus edificios. La estructura urbana seleccionada para este análisis comparativo es una red ortogonal con las calles principales orientadas este-oeste. El edificio típico que compone el vecindario se ha definido como “business as usual” (BAU) y se ha definido con una cubierta de baldosa de hormigón estándar, con un albedo 0.3, paredes con albedo 0.2 (construcción de muro de ladrillo típico) y cerramientos adiabáticos para evitar las posibles interferencias causadas por el intercambio térmico con el ambiente interior del edificio en los resultados del análisis. Para el caso de la cubierta ajardinada, se mantiene la misma geometría y características del edificio con excepción de la cobertura superficial de la azotea. Las baldosas de hormigón se han modificado con una cubierta ajardinada extensiva cubierta con plantas xerófilas, típicas en el clima de Madrid y caracterizado por su índice de densidad foliar, el “leaf area density” (LAD), que es la superficie total de superficie de hojas por unidad de volumen (m2/m3). El análisis se centra en los cañones urbanos entendidos como el espacio de calle comprendido entre los límites geométricos de la calle, verticales y horizontales, y el nivel superior de la cota urbana nivel de cubiertas. Los escenarios analizados se basan en la variación de la los principales parámetros que según la literatura analizada condicionan las variaciones microclimáticas en el ámbito urbano afectado por la vegetación, la velocidad del viento y el LAD de la azotea. Los resultados han sido registrados bajo condiciones de exposición solar diferentes. Las simulaciones fueron realizadas por los patrones de viento típico de verano, que para Madrid se caracterizan por vientos de componente suroeste que van desde 3 a 0 m/s. las simulaciones fueron realizadas para unas condiciones climáticas de referencia de 3, 2, 1 y 0 m/s a nivel superior del cañón urbano, como condición de contorno para el análisis. Los resultados calculados a 1,4 metros por encima del nivel del suelo, en el espacio habitado, mostraron que el efecto de la cubierta ajardinada era menor en condiciones de contorno con velocidades de viento más altas aunque en ningún caso el efecto de la cubierta verde sobre la temperatura del aire superó reducciones de temperatura de aire superiores a 1 º C. La humedad relativa no presentó variaciones significativas al comparar los diferentes escenarios. Las simulaciones realizadas para vientos con velocidad baja, entre 0 y 1 m/s mostraron que por debajo de 0.5 m/s la turbulencia del modelo aumentó drásticamente y se convirtió en el modelo inestable e incapaz de producir resultados fiables. Esto es debido al modelo de turbulencia en el software que no es válido para velocidades de viento bajas, lo que limita la capacidad de ENVI-met 4 para realizar simulaciones en estas condiciones de viento y es una de las principales conclusiones de este análisis en cuanto a la herramienta de simulación. También se comprobó el efecto de las densidades de la densidad de hoja (LAD) de los componentes vegetales en el modelo en la capa de aire inmediatamente superior a la cubierta, a 0,5 m sobre este nivel. Se compararon tres alternativas de densidad de hoja con la cubierta de baldosa de hormigón: el techo verde con LAD 0.3 (hierba típica o sedum), LAD 1.5 (plantas mixtas típicas) y LAD 2.5 (masa del árbol). Los resultados mostraron diferencias de temperatura muy relevante entre las diferentes alternativas de LAD analizadas. Los resultados muestran variaciones de temperatura que oscilan entre 3 y 5 º C al comparar el estándar de la azotea concreta con albedo 0, 3 con el techo con vegetación y vegetación densa, mostrando la importancia del LAD en la cuantificación de los efectos de las cubiertas vegetales en microclima circundante, lo que coincide con los datos reportados en la literatura existente y con los estudios empíricos analizados. Los resultados de los análisis teóricos han llegado a las siguientes conclusiones iniciales relacionadas con la herramienta de simulación y los resultados del modelo: En relación con la herramienta ENVI-met, se han observado limitaciones para el análisis. En primer lugar, la estructura rígida de la geometría, las bases de datos y el tamaño de la cuadrícula, limitan la escala y resolución de los análisis no permitiendo el desarrollo de grandes zonas urbanas. Por otro lado la estructura de ENVI-met permite el desarrollo de este tipo de simulación tan complejo dentro de tiempos razonables de cálculo y requerimientos computacionales convencionales. Otra limitación es el modelo de turbulencia del software, que no modela correctamente velocidades de viento bajas (entre 0 y 1 m/s), por debajo de 0,5 m/s el modelo da errores y no es estable, los resultados a estas velocidades no son fiables porque las turbulencias generadas por el modelo hacen imposible la extracción de patrones claros de viento y temperatura que permitan la comparación entre los escenarios de cubierta de hormigón y ajardinada. Además de las limitaciones anteriores, las bases de datos y parámetros de entrada en la versión pública del software están limitados y la complejidad de generar nuevos sistemas adaptándolos al edificio o modelo urbano que se quiera reproducir no es factible salvo en la versión profesional del software. Aparte de las limitaciones anteriores, los patrones de viento y perfiles de temperatura generados por ENVI-met concuerdan con análisis previos en los que se identificaban patrones de variación de viento y temperaturas en cañones urbanos con patrones de viento, relación de aspecto y dimensiones similares a los analizados en esta investigación. Por lo tanto, el software ha demostrado una buena capacidad para reproducir los patrones de viento en los cañones de la calle y capturar el efecto de enfriamiento producido por la cubierta verde en el cañón. En relación con el modelo, el resultado revela la influencia del viento, la radiación y el LAD en la temperatura del aire en cañones urbanos con relación de aspecto comprendida entre 0,5 y 1. Siendo el efecto de la cubierta verde más notable en cañones urbanos sombreados con relación de aspecto 1 y velocidades de viento en el nivel de “canopy” (por encima de la cubierta) de 1 m/s. En ningún caso las reducciones en la temperatura del aire excedieron 1 º C, y las variaciones en la humedad relativa no excedieron 1% entre los escenarios estudiados. Una vez que se han identificado los parámetros relevantes, que fueron principalmente la velocidad del viento y el LAD, se realizó un análisis experimental para comprobar los resultados obtenidos por el modelo. Para éste propósito se identificó una cubierta ajardinada de grandes dimensiones capaz de representar la escala urbana que es el objeto del estudio. El edificio usado para este fin fue el parking de la terminal 4 del aeropuerto internacional de Madrid. Aunque esto no es un área urbana estándar, la escala y la configuración del espacio alrededor del edificio fueron considerados aceptables para el análisis por su similitud con el contexto urbano objeto de estudio. El edificio tiene 800 x 200 m, y una altura 15 m. Está rodeado de vías de acceso pavimentadas con aceras conformando un cañón urbano limitado por el edificio del parking, la calle y el edificio de la terminal T4. El aparcamiento está cerrado con fachadas que configuran un espacio urbano de tipo cañón, con una relación de aspecto menor que 0,5. Esta geometría presenta patrones de viento y velocidad dentro del cañón que difieren ligeramente de los generados en el estudio teórico y se acercan más a los valores a nivel de canopo sobre la cubierta del edificio, pero que no han afectado a la tendencia general de los resultados obtenidos. El edificio cuenta con la cubierta ajardinada más grande en Europa, 12 Ha cubiertas por con una mezcla de hierbas y sedum y con un valor estimado de LAD de 1,5. Los edificios están rodeados por áreas plantadas en las aceras y árboles de sombra en las fachadas del edificio principal. El efecto de la cubierta ajardinada se evaluó mediante el control de temperaturas y humedad relativa en el cañón en un día típico de verano. La selección del día se hizo teniendo en cuenta las predicciones meteorológicas para que fuesen lo más semejantes a las condiciones óptimas para capturar el efecto de la cubierta vegetal sobre el microclima urbano identificadas en el modelo teórico. El 09 de julio de 2014 fue seleccionado para la campaña de medición porque las predicciones mostraban 1 m/s velocidad del viento y cielos despejados, condiciones muy similares a las condiciones climáticas bajo las que el efecto de la cubierta ajardinada era más notorio en el modelo teórico. Las mediciones se registraron cada hora entre las 9:00 y las 19:00 en 09 de julio de 2014. Temperatura, humedad relativa y velocidad del viento se registraron en 5 niveles diferentes, a 1.5, 4.5, 7.5, 11.5 y 16 m por encima del suelo y a 0,5 m de distancia de la fachada del edificio. Las mediciones fueron tomadas en tres escenarios diferentes, con exposición soleada, exposición la sombra y exposición influenciada por los árboles cercanos y suelo húmedo. Temperatura, humedad relativa y velocidad del viento se registraron con un equipo TESTO 410-2 con una resolución de 0,1 ºC para temperatura, 0,1 m/s en la velocidad del viento y el 0,1% de humedad relativa. Se registraron las temperaturas de la superficie de los edificios circundantes para evaluar su efecto sobre los registros usando una cámara infrarroja FLIR E4, con resolución de temperatura 0,15ºC. Distancia mínima a la superficie de 0,5 m y rango de las mediciones de Tª de - 20 º C y 250 º C. Los perfiles de temperatura extraídos de la medición in situ mostraron la influencia de la exposición solar en las variaciones de temperatura a lo largo del día, así como la influencia del calor irradiado por las superficies que habían sido expuestas a la radiación solar así como la influencia de las áreas de jardín alrededor del edificio. Después de que las medidas fueran tomadas, se llevaron a cabo las siguientes simulaciones para evaluar el impacto de la cubierta ajardinada en el microclima: a. estándar de la azotea: edificio T4 asumiendo un techo de tejas de hormigón con albedo 0.3. b. b. cubierta vegetal : T4 edificio asumiendo una extensa cubierta verde con valor bajo del LAD (0.5)-techo de sedum simple. c. c. cubierta vegetal: T4 edificio asumiendo una extensa cubierta verde con alta joven valor 1.5-mezcla de plantas d. d. cubierta ajardinada más vegetación nivel calle: el edificio T4 con LAD 1.5, incluyendo los árboles existentes a nivel de calle. Este escenario representa las condiciones actuales del edificio medido. El viento de referencia a nivel de cubierta se fijó en 1 m/s, coincidente con el registro de velocidad de viento en ese nivel durante la campaña de medición. Esta velocidad del viento se mantuvo constante durante toda la campaña. Bajo las condiciones anteriores, los resultados de los modelos muestran un efecto moderado de azoteas verdes en el microclima circundante que van desde 1 º a 2 º C, pero una contribución mayor cuando se combina con vegetación a nivel peatonal. En este caso las reducciones de temperatura alcanzan hasta 4 ºC. La humedad relativa sin embargo, no presenta apenas variación entre los escenarios con y sin cubierta ajardinada. Las temperaturas medidas in situ se compararon con resultados del modelo, mostrando una gran similitud en los perfiles definidos en ambos casos. Esto demuestra la buena capacidad de ENVI-met para reproducir el efecto de la cubierta ajardinada sobre el microclima y por tanto para el fin de esta investigación. Las diferencias más grandes se registraron en las áreas cercanas a las zonas superiores de las fachadas que estaban más expuestas a la radiación del sol y también el nivel del suelo, por la influencia de los pavimentos. Estas diferencias se pudieron causar por las características de los cerramientos en el modelo que estaban limitados por los datos disponibles en la base de datos de software, y que se diferencian con los del edificio real. Una observación importante derivada de este estudio es la contribución del suelo húmedo en el efecto de la cubierta ajardinada en la temperatura del aire. En el escenario de la cubierta ajardinada con los arboles existentes a pie de calle, el efecto del suelo húmedo contribuye a aumentar las reducciones de temperatura hasta 4.5ºC, potenciando el efecto combinado de la cubierta ajardinada y la vegetación a pie de calle. Se realizó un análisis final después de extraer el perfil horario de temperaturas en el cañón urbano influenciado por el efecto de las cubiertas ajardinadas y los árboles. Con esos perfiles modificados de temperatura y humedad se desarrolló un modelo energético en el edificio asumiendo un edificio cerrado y climatizado, con uso de oficinas, una temperatura de consigna de acuerdo al RITE de 26 ºC, y con los sistemas por defecto que establece el software para el cálculo de la demanda energética y que responden a ASHRAE 90.1. El software seleccionado para la simulación fue Design Builder, por su capacidad para generar simulaciones horarias y por ser una de las herramientas de simulación energética más reconocidas en el mercado. Los perfiles modificados de temperatura y humedad se insertaron en el año climático tipo y se condujo la simulación horaria para el día definido, el 9 de Julio. Para la simulación se dejaron por defecto los valores de conductancia térmica de los cerramientos y la eficiencia de los equipos de acuerdo a los valores que fija el estándar ASHRAE para la zona climática de Madrid, que es la 4. El resultado mostraba reducciones en el consumo de un día pico de hasta un 14% de reducción en las horas punta. La principal conclusión de éste estudio es la confirmación del potencial de las cubiertas ajardinadas como una estrategia para reducir la temperatura del aire y consumo de energía en los edificios, aunque este efecto puede ser limitado por la influencia de los vientos, la radiación y la especie seleccionada para el ajardinamiento, en especial de su LAD. Así mismo, en combinación con los bosques urbanos su efecto se potencia e incluso más si hay pavimentos húmedos o suelos porosos incluidos en la morfología del cañón urbano, convirtiéndose en una estrategia potencial para adaptar los ecosistemas urbanos el efecto aumento de temperatura derivado del cambio climático. En cuanto a la herramienta, ENVI-met se considera una buena opción para éste tipo de análisis dada su capacidad para reproducir de un modo muy cercano a la realidad el efecto de las cubiertas. Aparte de ser una herramienta validada en estudios anteriores, en el caso experimental se ha comprobado por medio de la comparación de las mediciones con los resultados del modelo. A su vez, los resultados y patrones de vientos generados en los cañones urbanos coinciden con otros estudios similares, concluyendo por tanto que es un software adecuado para el objeto de esta tesis doctoral. Como líneas de investigación futura, sería necesario entender el efecto de la cubierta ajardinada en el microclima urbano en diferentes zonas climáticas, así como un mayor estudio de otras variables que no se han observado en este análisis, como la temperatura media radiante y los indicadores de confort. Así mismo, la evaluación de otros parámetros que afectan el microclima urbano tales como variables geométricas y propiedades superficiales debería ser analizada en profundidad para tener un resultado que cubra todas las variables que afectan el microclima en el cañón urbano. ABSTRACT Climate Change is posing an urgency in the development of strategies able not only to mitigate but also adapt to the effects that this global problem is evidencing around the world. Heat waves, flooding and severe draughts increase the vulnerability of population, and this is especially critical in urban settlements. This has been extensively studied over the past decades, addressed from different perspectives and ranging from the regional heat island analysis to the building scale. Its understanding requires physical and dimensional analysis of this broad phenomenon and a deep analysis of the factors and the strategies which can offset it. In the search of solutions to this problem, green infrastructure elements such as green roofs, walls and urban forests arise as strategies able provide multiple regulating ecosystem services to the urban environment able to cope with climate change effects. This includes storm water management, heat island effect control, and improvement of air and water quality. Over the last decade, multiple studies have been developed to evaluate and quantify the ecosystem services provided by green roofs, however, specific regulating services addressing urban microclimate and their impact on the urban dwellers have not been widely quantified. This research tries to contribute to fill this gap and analyzes the effects of green roofs and urban forests on urban microclimate at pedestrian level, quantifying its potential for regulating ambient temperature in hot season in Mediterranean –continental climates. The study is divided into a sequence of analysis where the critical factors affecting the performance of the green roof system on the microclimate are identified and the effects of the green roof is tested in a real case study. The first step has been the definition of the object of study, through the analysis and review of theoretical and empirical papers that investigate the effects of covers landscaped in the built environment, in the context of its use as a tool for adaptation and mitigation of the impact of climate change on cities and urban development. This literature review, reveals the great potential of the plant systems as a tool for passive design capable of improving the climatic and microclimatic conditions in the cities, as well as its positive impact on the energy performance of buildings, but also the need for further analysis at the street scale where climate, urban surfaces and materials, and vegetation converge. This analysis requires a methodology where the thermal buildings response, the variations in the patterns of wind and the interaction of the vegetation are integrated, so a quantitative analysis can help to define the most effective strategies to achieve liveable urban spaces and collaterally, , the improvement of the surrounding buildings energy performance. In this specific scale research is needed and should be customized to every climate, urban condition and nature based strategy. In this context, the main objective for this research was the quantitative assessment of the Green roof impact on the urban microclimate at a neighbourhood scale in summer conditions in Mediterranean- continental climates. For the achievement of this main objective, the following secondary objectives have been set: • Identify the numerical models and calculation tools able to capture the effect of the roof garden on the microclimate. • Identify the enhancing or limiting parameter affecting this effect. • Quantification of the impact of the microclimate created on the energy consumption of buildings surrounding the street canyon analysed. The main hypothesis behind this research and where the above objectives are funded on is as follows: "An extensive roof installed in medium height buildings favours the establishment of microclimates at the pedestrian level and reduces the temperatures in the urban environment where they are located." For the purpose of verifying the above hypothesis and achieving the proposed objectives the following methodology has been followed: - Definition of hypothesis and objectives - Definition of the scope and limitations - Theoretical analysis of parameters affecting gren roof performance - Experimental analysis; - Energy modelling analyisis - Conclusions and future lines of work The search for suitable tools and models for meeting the objectives of this analysis has led to ENVI-met v4 as the most suitable software for this research. ENVI met is a three-dimensional micro-climate model designed to simulate the surface-plant-air interactions in urban environments. Based in the fundamental equations representing, mass, energy and momentum conservation, the software has the capacity of representing the complex phenomena characterizing the microclimate in urban canyons, overcoming the challenge posed by the computing requirements of a full calculus based on finite elements done via traditional computational fluid dynamics tools. Once the analysis tool has been defined, a first set of analysis has been developed to identify the main parameters affecting the green roof influence on the microclimate. In this analysis, two different scenarios are compared. A neighborhood with standard concrete tile roof and the same configuration substituting the concrete tile by an extensive green roof. Once the scenarios have been modeled, different iterations have been run to identify the influence of different wind patterns, solar exposure and roof vegetation type on the microclimate, since those are the most relevant variables affecting urban microclimates. These analysis have been run to check the conditions under which the effects of green roofs get significance. Since ENVI-met V4 is a predictive software, the first step has been the definition of the initial weather conditions which are then used as starting point by the software, which generates its own daily temperature and humidity profile based on the location of the building, geometry, vegetation and the surfaces physical characteristics. The base geometry used for this first analysis has been based on a typical urban layout structure located in Madrid, an orthogonal net with the main streets oriented East-West to ease the analysis of solar radiation in the different points of the model. This layout represents a typical urban neighborhood, with street canyons keeping an aspect ratio between 0.5 and 1 and high sky view factor to ensure correct sun access to the streets and buildings and work with typical wind flow patterns. Finally, the roof vegetation has been defined in terms of foliage density known as Leaf Area Density (LAD) and defined as the total one-sided leaf area per unit of layer volume. This index is the most relevant vegetation characteristic for the purpose of calculating the effect of vegetation on wind and solar radiation as well as the energy consumed during its metabolic processes. The building as usual (BAU) configuring the urban layout has been defined with standard concrete tile roofs, considering 0.3 albedo. Walls have been set with albedo 0.2 (typical brick wall construction) and adiabatic to avoid interference caused by thermal interchanges with the building indoor environment. For the proposed case, the same geometry and building characteristics have been kept. The only change is the roof surface coverage. The gravel on the roof has been changed with an extensive green roof covered with drought tolerant plants, typical in Madrid climate, and characterized by their LAD. The different scenarios analysed are based in the variation of the wind speed and the LAD of the roof. The results have been recorded under different sun exposure conditions. Simulations were run for the typical summer wind patterns, that for Madrid are characterized by South-west winds ranging from 3 to 0 m/s. Simulations were run for 3, 2, 1 and 0 m/s at urban canopy level. Results taken at 1.4 m above the ground showed that the green roof effect was lower with higher wind speeds and in any case the effect of the green roof on the air temperatures exceeded air temperature reductions higher than 1ºC. Relative humidity presented no variations when comparing the different scenarios. For the analysis at 0m/s, ENVI-met generated error and no results were obtained. Different simulations showed that under 0.5 m/s turbulence increased dramatically and the model became unstable and unable to produce reliable results. This is due to the turbulence model embedded in the software which is not valid for low wind speeds (below 1 m/s). The effect of the different foliage densities was also tested in the model. Three different alternatives were compared against the concrete roof: green roof with LAD 0.3 ( typical grass or sedum), 1.5 (typical mixed plants) and 2.5 (tree mass). The results showed very relevant temperature differences between the different LAD alternatives analyzed. Results show temperature variations ranging between 3 and 5 ºC when comparing the standard concrete roof with albedo 0, 3 with the vegetated roof and vegetated mass, showing the relevance of the LAD on the effects of green roofs on microclimate. This matches the data reported in existing literature and empirical studies and confirms the relevance of the LAD in the roof effect on the surrounding microclimate. The results of the theoretical analysis have reached the following initial conclusions related to both, the simulation tool and the model results: • In relation to the tool ENVI-met, some limitations for the analysis have been observed. In first place, the rigid structure of the geometry, the data bases and the grid size, limit the scale and resolution of the analysis not allowing the development of large urban areas. On the other hand the ENVI-met structure enables the development of this type of complex simulation within reasonable times and computational requirements for the purpose of this analysis. Additionally, the model is unable to run simulations at wind speeds lower than 0.5 m/s, and even at this speed, the results are not reliable because the turbulences generated by the model that made impossible to extract clear temperature differences between the concrete and green roof scenarios. Besides the above limitations, the wind patterns and temperature profiles generated by ENVImet are in agreement with previous analysis identifying wind patterns in urban canyons with similar characteristics and aspect ratio. Therefore the software has shown a good capacity for reproducing the wind effects in the street canyons and seems to capture the cooling effect produced by the green roof. • In relation to the model, the results reveals the influence of wind, radiation and LAD on air temperature in urban canyons with aspect ratio comprised between 0.5 and 1. Being the effect of the green roof more noticeable in shaded urban canyons with aspect ratio 1 and wind speeds of 1 m/s. In no case the reductions in air temperature exceeded 1ºC. Once the relevant parameters have been identified, mainly wind speed and LAD, an experimental analysis was conducted to test the results obtained by the model. For this purpose a large green roof was identified, able to represent the urban scale which is the object of the studio. The building identified for this purpose was the terminal 4, parking building of the international Madrid Airport. Even though this is not a standard urban area, the scale and configuration of the space around the building were deemed as acceptable for the analysis. The building is an 800x200 m, 15 m height parking building, surrounded by access paved paths and the terminal building. The parking is enclosed with facades that configure an urban canyon-like space, although the aspect ratio is lower than 0.5 and the wind patterns might differ from the theoretical model run. The building features the largest green roof in Europe, a 12 Ha extensive green roof populated with a mix of herbs and sedum with a LAD of 1.5. The buildings are surrounded by planted areas at the sidewalk and trees shading the main building facades. Green roof performance was evaluated by monitoring temperatures and relative humidity in the canyon in a typical summer day. The day selection was done taking into account meteorological predictions so the weather conditions on the measurement day were as close as possible as the optimal conditions identified in terms of green roof effects on the urban canyon. July 9th 2014 was selected for the measurement campaign because the predictions showed 1 m/s wind speed and sunny sky, which were very similar to the weather conditions where the effect of the green roof was most noticeable in the theory model. Measurements were registered hourly from 9:00am to 19:00 on July 9th 2014. Temperature, relative humidity and wind speed were recorded at 5 different levels, at 1.5, 4.5, 7.5, 11.5 and 16 m above ground and at 0.5 m distance from the building façade. Measurements were taken in three different scenarios, sunny exposure, shaded exposure, and shaded exposure influenced by nearby trees and moist soil. Temperature, relative humidity and wind speed were registered using a TESTO 410-2 anemometer, with 0.1ºC resolution for temperature, 0.1 m/s resolution for wind speed and 0.1 % for relative humidity. Surface temperatures were registered using an infrared camera FLIR E4, with temperature resolution 0.15ºC. Minimal distance to surface of 0.5 m and Tª measurements range from -20ºC and 250ºC. The temperature profiles measured on the site showed the influence of solar exposure on the temperature variations along the day, as well as the influence of the heat irradiated by the building surfaces which had been exposed to the sun radiation and those influenced by the moist soft areas around the building. After the measurements were taken, the following simulations were conducted to evaluate the impact of the green roof on the microclimate: a. Standard roof: T4 building assuming a concrete tile roof with albedo 0.3. b. Green roof: T4 building assuming an extensive green roof with low LAD value (0.5)-Simple Sedum roof. c. Green roof: T4 building assuming an extensive green roof with high LAD value 1.5- Lucerne and grasses d. Green roof plus street level vegetation: T4 Building, LAD 1.5 (Lucerne), including the existing trees at street level. This scenario represents the current conditions of the building. The urban canopy wind was set as 1 m/s, the wind speed register at that level during the measurement campaign. This wind speed remained constant over the whole campaign. Under the above conditions, the results of the models show a moderate effect of green roofs on the surrounding microclimate ranging from 1ºC to 2ºC, but a larger contribution when combining it with vegetation at pedestrian level, where 4ºC temperature reductions are reached. Relative humidity remained constant. Measured temperatures and relative humidity were compared to model results, showing a close match in the profiles defined in both cases and the good capacity of ENVI met to capture the impact of the green roof in this analysis. The largest differences were registered in the areas close to the top areas of the facades which were more exposed to sun radiation and also near to the soil level. These differences might be caused by differences between the materials properties included in the model (which were limited by the data available in the software database) and those in the real building. An important observation derived from this study is the contribution of moist soil to the green roof effect on air temperatures. In the green roof scenario with surrounding trees, the effect of the moist soil contributes to raise the temperature reductions at 4.5ºC. A final analysis was conducted after extracting the hourly temperature profile in the street canyon influenced by the effect of green roofs and trees. An energy model was run on the building assuming it was a conventional enclosed building. Energy demand reductions were registered in the building reaching up to 14% reductions at the peak hour. The main conclusion of this study is the potential of the green roofs as a strategy for reducing air temperatures and energy consumption in the buildings, although this effect can be limited by the influence of high speed winds. This effect can be enhanced its combination with urban forests and even more if soft moist pavements are included in the urban canyon morphology, becoming a potential strategy for adapting urban ecosystems to the increasing temperature effect derived from climate change.

The specific features of methionine biosynthesis and metabolism in plants

Plants, unlike other higher eukaryotes, possess all the necessary enzymatic equipment for de novo synthesis of methionine, an amino acid that supports additional roles than simply serving as a building block for protein synthesis. This is because methionine is the immediate precursor of S-adenosylmethionine (AdoMet), which plays numerous roles of being the major methyl-group donor in transmethylation reactions and an intermediate in the biosynthesis of polyamines and of the phytohormone ethylene. In addition, AdoMet has regulatory function in plants behaving as an allosteric activator of threonine synthase. Among the AdoMet-dependent reactions occurring in plants, methylation of cytosine residues in DNA has raised recent interest because impediment of this function alters plant morphology and induces homeotic alterations in flower organs. Also, AdoMet metabolism seems somehow implicated in plant growth via an as yet fully understood link with plant-growth hormones such as cytokinins and auxin and in plant pathogen interactions. Because of this central role in cellular metabolism, a precise knowledge of the biosynthetic pathways that are responsible for homeostatic regulation of methionine and AdoMet in plants has practical implications, particularly in herbicide design.

Molecular Characterization of the Oxalate Oxidase Involved in the Response of Barley to the Powdery Mildew Fungus1

Previously we reported that oxalate oxidase activity increases in extracts of barley (Hordeum vulgare) leaves in response to the powdery mildew fungus (Blumeria [syn. Erysiphe] graminis f.sp. hordei) and proposed this as a source of H2O2 during plant-pathogen interactions. In this paper we show that the N terminus of the major pathogen-response oxalate oxidase has a high degree of sequence identity to previously characterized germin-like oxalate oxidases. Two cDNAs were isolated, pHvOxOa, which represents this major enzyme, and pHvOxOb', representing a closely related enzyme. Our data suggest the presence of only two oxalate oxidase genes in the barley genome, i.e. a gene encoding HvOxOa, which possibly exists in several copies, and a single-copy gene encoding HvOxOb. The use of 3′ end gene-specific probes has allowed us to demonstrate that the HvOxOa transcript accumulates to 6 times the level of the HvOxOb transcript in response to the powdery mildew fungus. The transcripts were detected in both compatible and incompatible interactions with a similar accumulation pattern. The oxalate oxidase is found exclusively in the leaf mesophyll, where it is cell wall located. A model for a signal transduction pathway in which oxalate oxidase plays a central role is proposed for the regulation of the hypersensitive response.

A Carboniferous insect gall: insight into early ecologic history of the Holometabola.

Although the prevalence or even occurrence of insect herbivory during the Late Carboniferous (Pennsylvanian) has been questioned, we present the earliest-known ecologic evidence showing that by Late Pennsylvanian times (302 million years ago) a larva of the Holometabola was galling the internal tissue of Psaronius tree-fern fronds. Several diagnostic cellular and histological features of these petiole galls have been preserved in exquisite detail, including an excavated axial lumen filled with fecal pellets and comminuted frass, plant-produced response tissue surrounding the lumen, and specificity by the larval herbivore for a particular host species and tissue type. Whereas most suggestions over-whelmingly support the evolution of such intimate and reciprocal plant-insect interactions 175 million years later, we provide documentation that before the demise of Pennsylvanian age coal-swamp forests, a highly stereotyped life cycle was already established between an insect that was consuming internal plant tissue and a vascular plant host responding to that herbivory. This and related discoveries of insect herbivore consumption of Psaronius tissues indicate that modern-style herbivores were established in Late Pennsylvanian coal-swamp forests.

Identifying functional domains within terpene cyclases using a domain-swapping strategy.

Cyclic terpenes and terpenoids are found throughout nature. They comprise an especially important class of compounds from plants that mediate plant- environment interactions, and they serve as pharmaceutical agents with antimicrobial and anti-tumor activities. Molecular comparisons of several terpene cyclases, the key enzymes responsible for the multistep cyclization of C10, C15, and C20 allylic diphosphate substrates, have revealed a striking level of sequence similarity and conservation of exon position and size within the genes. Functional domains responsible for a terminal enzymatic step were identified by swapping regions approximating exons between a Nicotiana tabacum 5-epi-aristolochene synthase (TEAS) gene and a Hyoscyamus muticus vetispiradiene synthase (HVS) gene and by characterization of the resulting chimeric enzymes expressed in bacteria. While exon 4 of the TEAS gene conferred specificity for the predominant reaction products of the tobacco enzyme, exon 6 of the HVS gene conferred specificity for the predominant reaction products of the Hyoscyamus enzyme. Combining these two functional domains of the TEAS and HVS genes resulted in a novel enzyme capable of synthesizing reaction products reflective of both parent enzymes. The relative ratio of the TEAS and HVS reaction products was also influenced by the source of exon 5 present in the new chimeric enzymes. The association of catalytic activities with conserved but separate exonic domains suggests a general means for generating additional novel terpene cyclases.

Interazioni piante - insetti: ruolo delle formiche in un sistema multitrofico. Aspetti ecoetologici ed applicativi

Le formiche svolgono un importante ruolo all’interno degli ecosistemi ed alcune specie sono considerate keystone in quanto in grado di modificare la componente biotica e/o abiotica dell’ecosistema stesso. Sono animali ubiquitari che hanno colonizzato molteplici ambienti, compresi gli agroecosistemi. Negli agroecosistemi spesso svolgono un ruolo impattante determinando la diffusione o il regresso di specie di artropodi, alcune delle quali dannose alle colture. La presente ricerca tiene conto di un’ampia visione dei rapporti ecoetologici intercorrenti tra le formiche e la componente biotica di un ecosistema, utilizzando il concetto di rete multitrofica. In quest’ottica, si è pensato di costruire un sistema multitrofico costituito da una specie vegetale di interesse agrario (Cucumis sativus), dai suoi fitofagi naturali, divisi in fitomizi (afidi) (Aphis gossypii e Myzus persicae) e fitofagi masticatori (bruchi del lepidottero Mamestra brassicae), formiche (Formica pratensis) e predatori afidofagi (Aphidolets aphidimyza). Il sistema multitrofico è stato utilizzato sia per studiare l’aggressività delle formiche, sia per verificare l’esistenza di una comunicazione interspecifica tra le formiche e le piante (allelochimici). Gli studi sull’aggressività sono consistiti nel: • Verificare il livello di aggressività delle formiche nei confronti di un fitofago masticatore, competitore degli afidi nello sfruttare la pianta ospite. • Verificare se la presenza di afidi mutualisti fa variare il livello di aggressività delle formiche verso il competitore. • Verificare se esiste aggressività verso un predatore di afidi, i quali, secondo il paradigma della trofobiosi, dovrebbero essere difesi dalle formiche in cambio della melata. • Verificare se il predatore ha evoluto strategie volte ad eludere il controllo delle formiche sugli insetti che si approcciano alla colonia di afidi. Gli studi sui rapporti piante-formiche sono stati effettuati mediante olfattometro, osservando la risposta delle formiche alle sostanze volatili provenienti da piante infestate in modo differente con i fitofagi del sistema. Attraverso il trappolaggio e l’analisi gas-cromatografica delle sostanze prodotte dalle piante oggetto di studio abbiamo quindi individuato tipo e quantità di ogni composto volatile. Oltre alle piante di cetriolo, per questi esperimenti sono state utilizzate anche piante di patata (Solanum tuberosum). Dagli esperimenti sull’aggressività è risultato che le formiche manifestano un elevato potenziale predatorio, eradicando completamente la presenza dei bruchi sulle piante. Questo livello di aggressività tuttavia non cresce con la presenza degli afidi mutualisti che dovrebbero essere difesi dai competitori. Le formiche inoltre non sono in grado di sopprimere i predatori afidofagi che ipotizziamo riescano ad effettuare un camuffamento chimico, assumendo gli odori degli afidi dei quali si nutrono. I risultati degli esperimenti in olfattometro mostrano una chiara risposta positiva delle formiche verso gli odori di alcune delle piante infestate. Vi sono delle differenze nella risposta in funzione della specie di fitofago presente e della specie di pianta utilizzata. Nei trattamenti in cui erano presenti le piante di C. sativus, gli esperimenti in olfattometro hanno mostrato che le formiche rispondono in modo significativo agli odori emessi dalle piante in cui vi era la presenza del fitofago masticatore M. brassicae, solo o in associazione con A. gossypii. La presenza dei soli afidi, sia mutualisti (A. gossypii) sia non mutualisti (M. persicae), non ha invece indotto una risposta significativa nelle formiche rispetto agli odori delle piante non infestate. Nei trattamenti in cui erano presenti le piante di S. tuberosum la scelta delle formiche è stata significativa verso gli odori emessi dalle piante infestate con ciascuna delle singole specie di erbivori rispetto alle piante non infestate. Gli esperimenti sull’analisi delle sostanze volatili emesse dalle piante hanno confermato che gli organismi vegetali sono una vera centrale di produzione biochimica, infatti ben 91 composti volatili diversi sono stati individuati dall’analisi gas-cromatografica delle piante di cetriolo e 85 in quelle di patata. Dalle elaborazioni effettuate, rispettivamente 27 e 4 di essi sono prodotti esclusivamente dalle piante attaccate dai fitofagi. In generale, il cambiamento più consistente è dato dalla quantità di alcune sostanze volatili emesse dalle piante infestate rispetto a quelle integre che determina un cambiamento nei rapporti tra le sostanze che compongono i volatiles. E’ probabile che l’effetto attrattivo esercitato sulle formiche sia dato da un Blend di sostanze più che dai singoli composti presenti

Floral structure, breeding system and fruit-set in the threatened sub-shrub Tetratheca juncea Smith (Tremandraceae)

Tetratheca juncea Smith (Tremandraceae) has undergone a range contraction of approx. 50 km in the last 100 years and is now listed as a vulnerable sub-shrub restricted to the central and north coast regions of New South Wales, Australia. There are approx. 250 populations in a 110 km north-south distribution and populations are usually small with fewer than 50 plants/clumps. The reproductive ecology of the species was studied to determine why seed-set is reportedly rare. Flowers are bisexual, odourless and nectarless. Flowers are presented dependentally and there are eight stamens recurved around the pistil. Anthers are poricidal, contain viable pollen and basally contain a deep-red tapetal fluid that is slightly oily. Thus flowers are presented for buzz pollinators, although none were observed at flowers during our study. The species was found to be facultatively xenogamous with only one in 50 glasshouse flowers setting seed autogamously, i.e. without pollinator assistance. Field studies revealed fertile fruit in 24 populations but production varied significantly across sites from exceedingly low (0.6 fruits per plant clump) to low (17 fruits per plant clump). Fruit-set ranged from 0 to 65%, suggesting that pollen vectors exist or that autogamy levels in the field are variable and higher than glasshouse results. Fruit production did not vary with population size, although in three of the five populations in the south-west region more than twice as much fruit was produced as in populations elsewhere. A moderately strong relationship between foliage volume and fruit : flower ratios suggests that bigger plants may be more attractive than smaller plants to pollinators. A review of Tetratheca pollination ecology revealed that several species are poorly fecund and pollinators are rare. The habitat requirements for Tetratheca, a genus of many rare and threatened species, is discussed. (C) 2003 Annals of Botany Company.

Coral reefs in a century of rapid environmental change

Coral reefs are the most diverse marine ecosystem and embrace possibly millions of plant, animal and protist species. Mutualistic symbioses are a fundamental feature of coral reefs that have been used to explain their structure, biodiversity and existence. Complex inter-relationships between hosts, habitats and symbionts belie closely coupled nutrient and community dynamics that create the circumstances for something from nothing (or the oasis in a nutrient desert). The flip side of these dynamics is a close dependency between species, which results in a series of non-linear relationships as conditions change. These responses are being highlighted as anthropogenic influences increase across the world's tropical and subtropical coastlines. Caribbean as well as Indo-Pacific coral populations are now in a serious decline in many parts of the world. This has resulted in a significant reorganization of how coral reef ecosystems function. Among the spectra of changes brought about by humans is rapid climate change. Mass coral bleaching - the loss of the dinoflagellate symbionts from reef-building corals - and mortality has affected the world's coral reefs with increasing frequency and intensity since the late 1970s. Mass bleaching events, which often cover thousands of square kilometres of coral reefs, are triggered by small increases (+1-3degreesC) in water temperature. These increases in sea temperature are often seen during warm phase weather conditions (e.g. ENSO) and are increasing in size and magnitude. The loss of living coral cover (e.g. 16% globally in 1998, an exceptionally warm year) is resulting in an as yet unspecified reduction in the abundance of a myriad of other species. Projections from general circulation models (GCM) used to project changes in global temperature indicate that conditions even under the mildest greenhouse gas emission scenarios may exceed the thermal tolerances of most reef-building coral communities. Research must now explore key issues such as the extent to which the thermal tolerances of corals and their symbionts are dynamic if bleaching and disease are linked; how the loss of high densities of reef-building coral will affect other dependent species; and, how the loss of coral populations will affect the millions of people globally who depend on coral reefs for their daily survival.

Discrimination among host plants (Leucaena species and accessions) by the psyllid pest Heteropsylla cubana and implications for understanding resistance

1 The herbivorous bug Heteropsylla cubana Crawford (Homoptera: Psyllidae) is a pest of the cattle fodder crop Leucaena (Leguminosae: Mimosoideae). The interaction between the psyllid and three varieties of its Leucaena host plant was investigated in relation to the apparent resistance of some Leucaena varieties (Leucaena leucocephala, Leucaena pallida and their hybrids) to attack. 2 Field trials demonstrated that adult psyllids distinguished among the different varieties of Leucaena over a distance, and were attracted to L. leucocephala in significantly higher numbers than to L. pallida or to the hybrid. Pesticide treatment increased the attractiveness of Leucaena plants, even of those deemed to be psyllid resistant. Numbers of psyllid eggs and nymphs, sampled in the field, reflect the arrival rates of adults at the three plant varieties. 3 Wavelength reflectance data of the three Leucaena varieties were not significantly different from one another, suggesting that psyllids cannot discriminate among the three plants using brightness or wavelength cues. There was a differential release of caryophyllene among the three varieties. Release of caryophyllene in L. leucocephala and the hybrid appeared to be influenced by environmental conditions. 4 Experiments demonstrated that caryophyllene (at least on its own) did not influence the behaviour of leucaena psyllids in relation to leucaena plants. 5 The results suggest that host plant volatiles cannot be dismissed as significant in the interaction between the leucaena psyllid and its Leucaena host plants. Further avenues for investigation are recommended and these are related to novel ways of understanding resistance in insect plant inter-relationships.

Rhizodeposits of Trifolium pratense and Lolium perenne: Their comparative effects on 2,4-D mineralization in two contrasting soils

Rhizosphere enhanced biodegradation of organic pollutants has been reported frequently and a stimulatory role for specific components of rhizodeposits postulated. As rhizodeposit composition is a function of plant species and soil type, we compared the effect of Lolium perenne and Trifolium pratense grown in two different soils (a sandy silt loam: pH 4, 2.8% OC, no previous 2,4-D exposure and a silt loam: pH 6.5, 4.3% OC, previous 2,4-D exposure) on the mineralization of the herbicide 2,4-D (2,4-dichlorophenoxyacetic acid). We investigated the relationship of mineralization kinetics to dehydrogenase activity, most probable number of 2,4-D degraders (MPN2,4-D) and 2,4-D degrader composition (using sequence analysis of the gene encoding alpha-ketoglutarate/2,4-D dioxygenase (tfdA)). There were significant (P < 0.01) plant-soil interaction effects on MPN2,4-D and 2,4-D mineralization kinetics (e.g. T pratense rhizodeposits enhanced the maximum mineralization rate by 30% in the acid sandy silt loam soil, but not in the neutral silt loam soil). Differences in mineralization kinetics could not be ascribed to 2,4-D degrader composition as both soils had tfdA sequences which clustered with tfdAs representative of two distinct classes of 2,4-D degrader: canonical R. eutropha JMP134-like and oligotrophic alpha-proteobacterial-like. Other explanations for the differential rhizodeposit effect between soils and plants (e.g. nutrient competition effects) are discussed. Our findings stress that complexity of soil-plant-microbe interactions in the rhizosphere make the occurrence and extent of rhizosphere-enhanced xenobiotic degradation difficult to predict.

Rhizodeposition and the enhanced mineralization of 2,4-dichlorophenoxyacetic acid in soil from the Trifolium pratense rhizosphere

Enhanced biodegradation of organic xenobiotic compounds in the rhizosphere is frequently recorded although the specific mechanisms are poorly understood. We have shown that the mineralization of 2,4-dichlorophenoxyacetic acid (2,4-D) is enhanced in soil collected from the rhizosphere of Trifolium pratense[e.g. maximum mineralization rate = 7.9 days(-1) and time at maximum rate (t(1)) = 16.7 days for 12-day-old T. pratense soil in comparison with 4.7 days(-1) and 25.4 days, respectively, for non-planted controls). The purpose of this study was to gain a better understanding of the plant-microbe interactions involved in rhizosphere-enhanced biodegradation by narrowing down the identity of the T. pratense rhizodeposit responsible for stimulating the microbial mineralization of 2,4-D. Specifically, we investigated the distribution of the stimulatory component(s) among rhizodeposit fractions (exudates or root debris) and the influence of soil properties and plant species on its production. Production of the stimulatory rhizodeposit was dependent on soil pH (e.g. t(1) for roots grown at pH 6.5 was significantly lower than for those grown at pH 4.4) but independent of soil inorganic N concentration. Most strikingly, the stimulatory rhizodeposit was only produced by T. pratense grown in non-sterile soil and was present in both exudates and root debris. Comparison of the effect of root debris from plant species (three each) from the classes monocotyledon, dicotyledon (non-legume) and dicotyledon (legume) revealed that legumes had by far the greatest positive impact on 2,4-D mineralization kinetics. We discuss the significance of these findings with respect to legume-rhizobia interactions in the rhizosphere.

Ecology of Leptocoris Hahn (Hemiptera: Rhopalidae) Soapberry Bugs in Australia

Soapberry bugs are worldwide seed predators of plants in the family Sapindaceae. Australian sapinds are diverse and widespread, consisting of about 200 native trees and shrubs. This flora also includes two introduced environmental weeds, plus cultivated lychee (Litchi chinensis Sonn.), longan (Dimocarpus longan Lour.) and rambutan (Nephelium lappaceum L.). Accordingly, Australian soapberry bugs may be significant in ecology, conservation and agriculture. Here we provide the first account of their ecology. We find five species of Leptocoris Hahn in Australia, and list sapinds that do and do not serve as reproductive hosts. From museum and field records we map the continental distributions of the insects and primary hosts. Frequency of occupation varies among host species, and the number of hosts varies among the insects. In addition, differences in body size and beak length are related to host use. For example, the long-beaked Leptocoris tagalicus Burmeister is highly polyphagous in eastern rainforests, where it occurs on at least 10 native and non-native hosts. It aggregates on hosts with immature fruit and commences feeding before fruits dehisce. Most of its continental range, however, matches that of a single dryland tree, Atalaya hemiglauca F. Muell., which has comparatively unprotected seeds. The taxon includes a smaller and shorter-beaked form that is closely associated with Atalaya, and appears to be taxonomically distinct. The other widespread soapberry bug is the endemic Leptocoris mitellatus Bergroth. It too is short-beaked, and colonises hosts phenologically later than L. tagalicus, as seeds become more accessible in open capsules. Continentally its distribution is more southerly and corresponds mainly to that of Alectryon oleifolius Desf. Among all host species, the non-native environmental weeds Cardiospermum L. and Koelreuteria Laxm. are most consistently attacked, principally by L. tagalicus. These recent host shifts have biocontrol implications. In contrast, the sapinds planted as fruit crops appear to be less frequently used at present and mainly by the longer-beaked species.