Functional response of Ulmus minor Mill. to drought, flooding and dutch elm disease

Ulmus minor es una especie arbórea originaria de Europa cuyas poblaciones han sido diezmadas por el hongo patógeno causante de la enfermedad de la grafiosis. La conservación de los olmos exige plantearse su propagación a través de plantaciones y conocer mejor su ecología y biología. Ulmus minor es un árbol de ribera, pero frecuentemente se encuentra alejado del cauce de arroyos y ríos, donde la capa freática sufre fuertes oscilaciones. Por ello, nuestra hipótesis general es que esta especie es moderadamente resistente tanto a la inundación como a la sequía. El principal objetivo de esta tesis doctoral es entender desde un punto de vista funcional la respuesta de U. minor a la inundación, la sequía y la infección por O. novo-ulmi; los factores que posiblemente más influyen en la distribución actual de U. minor. Con este objetivo se persigue dar continuidad a los esfuerzos de conservación de esta especie que desde hace años se dedican en varios centros de investigación a nivel mundial, ya que, entender mejor los mecanismos que contribuyen a la resistencia de U. minor ante la inoculación con O. novo-ulmi y factores de estrés abiótico ayudará en la selección y propagación de genotipos resistentes a la grafiosis. Se han planteado tres experimentos en este sentido. Primero, se ha comparado la tolerancia de brinzales de U. minor y U. laevis – otro olmo ibérico – a una inmersión controlada con el fin de evaluar su tolerancia a la inundación y comprender los mecanismos de aclimatación. Segundo, se ha comparado la tolerancia de brinzales de U. minor y Quercus ilex – una especie típica de ambientes Mediterránea secos – a la falta de agua en el suelo con el fin de evaluar el grado de tolerancia y los mecanismos de aclimatación a la sequía. El hecho de comparar dos especies contrastadas responde al interés en entender mejor cuales son los procesos que conducen a la muerte de una planta en condiciones de sequía – asunto sobre el que hay una interesante discusión desde hace algunos años. En tercer lugar, con el fin de entender mejor la resistencia de algunos genotipos de U. minor a la grafiosis, se han estudiado las diferencias fisiológicas y químicas constitutivas e inducidas por O. novo-ulmi entre clones de U. minor seleccionados a priori por su variable grado de resistencia a esta enfermedad. En el primer experimento se observó que los brinzales de U. minor sobrevivieron 60 días inmersos en una piscina con agua no estancada hasta una altura de 2-3 cm por encima del cuello de la raíz. A los 60 días, los brinzales de U. laevis se sacaron de la piscina y, a lo largo de las siguientes semanas, fueron capaces de recuperar las funciones fisiológicas que habían sido alteradas anteriormente. La conductividad hidráulica de las raíces y la tasa de asimilación de CO2 neta disminuyeron en ambas especies. Por el contrario, la tasa de respiración de hojas, tallos y raíces aumentó en las primeras semanas de la inundación, posiblemente en relación al aumento de energía necesario para desarrollar mecanismos de aclimatación a la inundación, como la hipertrofia de las lenticelas que se observó en ambas especies. Por ello, el desequilibrio del balance de carbono de la planta podría ser un factor relevante en la mortalidad de las plantas ante inundaciones prolongadas. Las plantas de U. minor (cultivadas en envases de 16 litros a media sombra) sobrevivieron por un prolongado periodo de tiempo en verano sin riego; la mitad de las plantas murieron tras 90 días sin riego. El cierre de los estomas y la pérdida de hojas contribuyeron a ralentizar las pérdidas de agua y tolerar la sequía en U. minor. Las obvias diferencias en tolerancia a la sequía con respecto a Q. ilex se reflejaron en la distinta capacidad para ralentizar la aparición del estrés hídrico tras dejar de regar y para transportar agua en condiciones de elevada tensión en el xilema. Más relevante es que las plantas con evidentes síntomas de decaimiento previo a su muerte exhibieron pérdidas de conductividad hidráulica en las raíces del 80% en ambas especies, mientras que las reservas de carbohidratos apenas variaron y lo hicieron de forma desigual en ambas especies. Árboles de U. minor de 5 y 6 años de edad (plantados en eras con riego mantenido) exhibieron una respuesta a la inoculación con O. novo-ulmi consistente con ensayos previos de resistencia. La conductividad hidráulica del tallo, el potencial hídrico foliar y la tasa de asimilación de CO2 neta disminuyeron significativamente en relación a árboles inoculados con agua, pero solo en los clones susceptibles. Este hecho enlaza con el perfil químico “más defensivo” de los clones resistentes, es decir, con los mayores niveles de suberina, ácidos grasos y compuestos fenólicos en estos clones que en los susceptibles. Ello podría restringir la propagación del hongo en el árbol y preservar el comportamiento fisiológico de los clones resistentes al inocularlos con el patógeno. Los datos indican una respuesta fisiológica común de U. minor a la inundación, la sequía y la infección por O. novo-ulmi: pérdida de conductividad hidráulica, estrés hídrico y pérdida de ganancia neta de carbono. Pese a ello, U. minor desarrolla varios mecanismos que le confieren una capacidad moderada para vivir en suelos temporalmente anegados o secos. Por otro lado, el perfil químico es un factor relevante en la resistencia de ciertos genotipos a la grafiosis. Futuros estudios deberían examinar como este perfil químico y la resistencia a la grafiosis se ven alteradas por el estrés abiótico. ABSTRACT Ulmus minor is a native European elm species whose populations have been decimated by the Dutch elm disease (DED). An active conservation of this species requires large-scale plantations and a better understanding of its biology and ecology. U. minor generally grows close to water channels. However, of the Iberian riparian tree species, U. minor is the one that spread farther away from rivers and streams. For these reasons, we hypothesize that this species is moderately tolerant to both flooding and drought stresses. The main aim of the present PhD thesis is to better understand the functional response of U. minor to the abiotic stresses – flooding and drought – and the biotic stress – DED – that can be most influential on its distribution. The overarching goal is to aid in the conservation of this emblematic species through a better understanding of the mechanisms that contribute to resistance to abiotic and biotic stresses; an information that can help in the selection of resistant genotypes and their expansion in large-scale plantations. To this end, three experiments were set up. First, we compared the tolerance to experimental immersion between seedlings of U. minor and U. laevis – another European riparian elm species – in order to assess their degree of tolerance and understand the mechanisms of acclimation to this stress. Second, we investigated the tolerance to drought of U. minor seedlings in comparison with Quercus ilex (an oak species typical of dry Mediterranean habitats). Besides assessing and understanding U. minor tolerance to drought at the seedling stage, the aim was to shed light into the functional alterations that trigger drought-induced plant mortality – a matter of controversy in the last years. Third, we studied constitutive and induced physiological and biochemical differences among clones of variable DED resistance, before and following inoculation with Ophiostoma novo-ulmi. The goal is to shed light into the factors of DED resistance that is evident in some genotypes of U. minor, but not others. Potted seedlings of U. minor survived for 60 days immersed in a pool with running water to approximately 2-3 cm above the stem collar. By this time, U. minor seedlings died, whereas U. laevis seedlings moved out of the pool were able to recover most physiological functions that had been altered by flooding. For example, root hydraulic conductivity and leaf photosynthetic CO2 uptake decreased in both species; while respiration initially increased with flooding in leaves, stems and roots possibly to respond to energy demands associated to mechanisms of acclimation to soil oxygen deficiency; as example, a remarkable hypertrophy of lenticels was soon observed in flooded seedlings of both species. Therefore, the inability to maintain a positive carbon balance somehow compromises seedling survival under flooding, earlier in U. minor than U. laevis, partly explaining their differential habitats. Potted seedlings of U. minor survived for a remarkable long time without irrigation – half of plants dying only after 90 days of no irrigation in conditions of high vapour pressure deficit typical of summer. Some mechanisms that contributed to tolerate drought were leaf shedding and stomata closure, which reduced water loss and the risk of xylem cavitation. Obviously, U. minor was less tolerant to drought than Q. ilex, differences in drought tolerance resulting mostly from the distinct capacity to postpone water stress and conduct water under high xylem tension among species. More relevant was that plants of both species exhibited similar symptoms of root hydraulic failure (i.e. approximately 80% loss of hydraulic conductivity), but a slight and variable depletion of non-structural carbohydrate reserves preceding dieback. Five- and six-year-old trees of U. minor (planted in the field with supplementary watering) belonging to clones of contrasted susceptibility to DED exhibited a different physiological response to inoculation with O. novo-ulmi. Stem hydraulic conductivity, leaf water potential and photosynthetic CO2 uptake decreased significantly relative to control trees inoculated with water only in DED susceptible clones. This is consistent with the “more defensive” chemical profile observed in resistant clones, i.e. with higher levels of saturated hydrocarbons (suberin and fatty acids) and phenolic compounds than in susceptible clones. These compounds could restrict the spread of O. novo-ulmi and contribute to preserving the near-normal physiological function of resistant trees when exposed to the pathogen. These results evidence common physiological responses of U. minor to flooding, drought and pathogen infection leading to xylem water disruption, leaf water stress and reduced net carbon gain. Still, seedlings of U. minor develop various mechanisms of acclimation to abiotic stresses that can play a role in surviving moderate periods of flood and drought. The chemical profile appears to be an important factor for the resistance of some genotypes of U. minor to DED. How abiotic stresses such as flooding and drought affect the capacity of resistant U. minor clones to face O. novo-ulmi is a key question that must be contemplated in future research.

Comparison and Assessment of Two Emission inventories for the Madrid Region

Emission inventories are databases that aim to describe the polluting activities that occur across a certain geographic domain. According to the spatial scale, the availability of information will vary as well as the applied assumptions, which will strongly influence its quality, accuracy and representativeness. This study compared and contrasted two emission inventories describing the Greater Madrid Region (GMR) under an air quality simulation approach. The chosen inventories were the National Emissions Inventory (NEI) and the Regional Emissions Inventory of the Greater Madrid Region (REI). Both of them were used to feed air quality simulations with the CMAQ modelling system, and the results were compared with observations from the air quality monitoring network in the modelled domain. Through the application of statistical tools, the analysis of emissions at cell level and cell – expansion procedures, it was observed that the National Inventory showed better results for describing on – road traffic activities and agriculture, SNAP07 and SNAP10. The accurate description of activities, the good characterization of the vehicle fleet and the correct use of traffic emission factors were the main causes of such a good correlation. On the other hand, the Regional Inventory showed better descriptions for non – industrial combustion (SNAP02) and industrial activities (SNAP03). It incorporated realistic emission factors, a reasonable fuel mix and it drew upon local information sources to describe these activities, while NEI relied on surrogation and national datasets which leaded to a poorer representation. Off – road transportation (SNAP08) was similarly described by both inventories, while the rest of the SNAP activities showed a marginal contribution to the overall emissions.

Optimizing rabbit production under heat stress: cage density and cage size during fattening, and breeding system and water quality on rabbit does

The aim of this work was to evaluate different management strategies to optimize rabbit production under chronic heat stress. To achieve it, three trials were conducted. In the first trial, to find the optimal cage density in tropical very dry forest condition, were measured growth performance, mortality rate, injured animals and carcass performance over an initial population of 300 cross-breed rabbits of New Zealand, California, Butterfly, Dutch and Satin, weaned at 30 days (535 ± 8 g, standard error). Treatments evaluated were: 6, 12, 18 and 24 rabbits/m2 (3, 6, 9 and 12 rabbits/cage, respectively, each cage of 0.5 m2). The maximal temperature-humidity index indicated a severe heat stress from weaning to 2.2 kg body weight (experimental time). At the end of experimental period 10, 20, 30 and 30 rabbits from the treatments of 6, 12, 18 and 24 rabbits/m2, respectively, were slaughtered and carcass performance recorded. Average daily gain and feed intake decreased by 0.31 ± 0.070 and 1.20 ± 0.25 g, respectively, per each unit that the density increased at the beginning of the experiment (P = 0.001). It increased the length of the fattening period by 0.91 ± 0.16 d (P = 0.001) per each unit of increment of density. However, rabbit production (kg/m2) increased linear and quadratically with the density (P < 0.008). Animals housed at the highest density compared to the lower one tended to show a higher incidence of ringworm (68.9 vs 39.4%; P = 0.075), injured animals (16.8 vs 3.03%; P = 0.12) and mortality (20.5 vs 9.63%; P = 0.043). The proportion of scapular fat (P = 0.042) increased linearly with increasing levels of density. Increasing density reduced linearly dorsal length (P = 0.001), and reduced linear and quadratically drip loss percentage (P = 0.097 and 0.018, respectively). In the second trial, 46 nulliparous rabbit does (23 clipped and 23 unclipped) with a BW of 3.67 ± 0.05 kg (s.e.) were used to evaluate heat stress and circadian rhythms comparing unclipped and clipped rabbit does, and to study if a more extensive breeding system increase litters performance at weaning without impairing rabbit doe performance,. Rectal temperature, feed and water 4 intake were recorded for 24 h. Rabbit does were mated 7 d after circadian measurements, and randomly assigned to two breeding systems. Control (C): mated at 14 d after parturition + litter weaned at 35 d of age. Extensive (E): mate at 21 after parturition + litter weaned at 42 d of age. The first three cycles were evaluated concerning to rabbit doe and litter performance. Two hundred twenty eight weaned rabbits, were divided into two cage sizes: 0.5 and 0.25 m2 with same density (16 rabbit/m2) and growing performance was recorded. Farm and rectal temperatures were minimal and feed and water intake maximal during the night (P < 0.001). Unclipped rabbit does showed higher rectal temperature (P = 0.045) and lower feed intake respect to clipped does (P = 0.019) which suggest a lower heat stress in the latter. Kits weaned per litter was reduced by 33% (P=0.038) in C group. This reduction was more important in the 2nd and 3rd cycles compared to the first (P ≤ 0.054). Rabbit doe feed efficiency tended to decrease in E respect C group (P = 0.093), whereas it was impaired from the first to the third cycle by 48% (P = 0.014). Growing rabbits from the E group were heavier at weaning (by 38%. P < 0.001), showed a higher feed intake (+7.4%) and lower feed efficiency (-8.4%) throughout the fattening period (P ≤ 0.056) respect to C group. Cage size had minor influence in growing performance. In the third trial, forty five non pregnant and non lactating rabbit does (21 nulliparous and 24 multiparous) were assigned randomly to farm water and to potable water to study if a water quality improvement can affect positively rabbit doe response to heat stress during pregnancy and lactation. A transponder was implanted in each animal to record subcutaneous temperature at 07:30 and 14:30 h. Experimental period extended from pregnancy (with no lactation) to the next lactation (until day 28). Body temperature and milk production were recorded daily, and body condition, feed and water intake weekly. Water quality did not affect any trait (P ≥ 0.15). Pregnant rabbit does were classified as does that weaned (W: 47%), not weaned (NW: 44%) or those pregnant that did not deliver (NB: 9%). Body temperature and feed intake decreased during pregnancy (P ≤ 0.031), but water intake remained constant. In this period body temperature decreased with metabolic weight (P ≤ 0.009). In W and NW does, 5 from mating to birth energy and protein balance impaired (P≤0.011). Body temperature of W does tended to be the lowest (P ≤ 0.090). Pregnancy length and total number of kits born tended to be longer and higher in NW than in W does (P = 0.10 and 0.053, respectively). Kit mortality at birth and from birth to 14 d of lactation was high, being worse for NW than for W does (97 vs. 40%; P<0.001). Body temperature during lactation was maximal at day 12, and milk production increased it (P ≤ 0.025). . In conclusion, in our heat stress conditions densities higher than 18 rabbits/m2 (34 kg/m2) at the end of fattening, are not recommended despite cage size, gestation and lactation productivity impaired not only when lactation is extended and along successive reproductive cycles but also due to a reduced embryo/kit survival and finally water quality improvement did not attenuate negative effect of heat stress. RESUMEN El propósito de éste trabajo fue evaluar diferentes estrategias de manejo para optimizar la producción de conejos bajo estrés térmico. Para lo cual se desarrollaron tres experimentos. En el primer experimento, para encontrar el número óptimo de gazapos por m2 de jaula durante el cebo en condiciones de bosque muy seco tropical, se estudiaron los rendimientos durante el cebo, mortalidad, animales lesionados y rendimiento de la canal sobre una población inicial de 300 conejos mestizos de Nueva Zelanda, California, Mariposa, Holandés y Satin, destetados a los 30 días de edad (535 ± 8g, error estándar). Los tratamientos evaluados fueron: 6, 12, 18 y 24 conejos/m2 (3, 6, 9 y 12 conejos/jaula, respectivamente, en jaulas de 0.5 m2). Durante el período experimental (destete a 2.2 kg de peso vivo), se observaron valores de THI correspondientes con un estrés térmico severo (THI max. De 31 a 35). Al final del período experimental, 10, 20, 30, y 30 conejos de los tratamientos con densidades de 6, 12, 18 y 24 conejos/m2, respectivamente, fueron sacrificados y su canal fue valorada. El promedio de la ganancia diaria y el consumo de alimento disminuyeron en 0.31 ± 0.070 y 1.20 ± 0.25 g, respectivamente, por cada unidad de incremento en la densidad al inicio del experimento (P=0.001). Esto alargó el período de engorde en 0.91 ± 0.16 d (P=0.001) por cada unidad de incremento de la densidad. Sin embargo, la producción de conejos (kg/m2) aumentó lineal y cuadráticamente con la densidad (P<0.008). Los animales alojados en las mayores densidades en comparación con el resto tendieron a mostrar una mayore incidencia de tiña (68.9 vs 39.4%; P=0.075), de cantidad de animales heridos (16.8 vs 3.03%; P=0.12), así como de mortalidad (20.5 vs 9.63%; P=0.043). El aumento en la densidad aumentó linealmente la proporción de grasa escapular (P=0.042) y redujo linealmente la longitud dorsal (P=0.001), y lineal y cuadráticamente el porcentaje de pérdida por goteo (P=0.018). En el segundo experimento, 46 conejas nulliparas (23 rasuradas y 23 no rasuradas) con un peso vivo de 3.67 ± 0.05 kg (e.e.) fueron usadas para evaluar el estrés 8 térmico y los ritmos circadianos comparando conejas rasuradas o no, y estudiar si un sistema de crianza más extensivo mejora el desempeño de la camada al destete sin perjudicar la productividad de la coneja. Durante 24 h se midió la temperatura rectal, consumo de alimento y de agua. Las conejas fueron montadas 7 días después, y distribuidas en dos sistemas de crianza. El control (C): monta a 14 días posparto y destete a 35 d de edad. El extensivo (E): monta a 21 días posparto y destete a 42 d de edad. Se controló la productividad de la coneja y la camada durante los tres primeros ciclos. Doscientos veintiocho gazapos fueron distribuidos en dos tamaños de jaulas (0.5 y 0.25 m2) con la misma densidad (16 conejos/m2) y se controlaron sus rendimientos productivos. Durante la noche se observaron los valores mínimos para la temperatura ambiental y rectal, y los máximos para consumo de alimento y agua (P< 0.001). Las conejas no rasuradas mostraron mayor temperatura rectal (P=0.045) y menores valores de consumo de alimento con respecto a las conejas rasuradas (P=0.019), lo que sugiere un menor estrés térmico en las últimas. El número de gazapos destetados por camada se redujo en 33% (P=0.038) en el grupo C. Este comportamiento se acentuó en el 2do y 3er ciclo en comparación con el primero (P≤0.054). La eficiencia alimenticia de las conejas tendió a disminuir en el grupo E con respecto al grupo C (P=0.093), dicha tendencia se acentúa del primer al tercer ciclo en un 48% (P=0.014). Los gazapos en fase de crecimiento provenientes del grupo E fueron más pesados al momento del destete (en 38% P<0.001), mostrando un mayor consumo de alimento (+7.4%) y menor eficiencia alimenticia (-8.4%) a lo largo del engorde (P≤0.056) con respecto al grupo C. El tamaño de la jaula tuvo una mínima influencia en el comportamiento durante el crecimiento de éstos gazapos. En el tercer experimento, cuarenta y cinco conejas no gestantes ni lactantes (21 nulíparas y 24 multíparas) se les asignó al azar agua dos tipos de agua: común de la granja y agua potable, con el fin de estudiar si una mejora en la calidad del agua puede afectar positivamente la respuesta de la coneja al estrés térmico durante la gestación y la lactancia. Se les implantó un transponder para registrar la temperatura subcutánea a las 7:30 y a las 14:30 h. El período experimental se extendió desde la gestación (sin 9 lactancia) hasta la lactanción consecutiva (hasta los 28 días). La temperatura corporal y la producción de leche se controlaron diariamente, y la condición corporal, consumo de agua y alimento, semanalmente. La calidad del agua no afectó a ninguna variable (P≥0.15). Las conejas preñadas fueron clasificadas como conejas que destetaron (W: 47%), que no destetaron (NW:44%) o aquellas que no parieron (NB: 9%). La temperatura corporal y consumo de alimento disminuyeron durante la gestación (P≤0.031), mientras que el consumo de agua se mantuvo constante. La temperatura corporal descendió con el peso metabólico durante la gestación (P≤0.009). El balance de energía y proteína disminuyó desde la monta al parto para las conejas W y NW (P≤0.011). Durante la gestación la temperatura corporal tendió a ser menor en las conejas W (P≤0.090). La longitud de la gestación y el número total de gazapos nacidos tendieron a ser mayores en conejas NW que en conejas W (P=0.10 y 0.053, respectivamente). La mortalidad de los gazapos al parto y del parto a los 14 días de lactancia fue alta, siendo peor para las conejas NW que para las W (97 vs 40%; P<0.001). Durante la lactancia la temperatura corporal alcanzó su valor máximo para el día 12, y la producción de leche indujo un incremento en la misma (P≤0.025). En conclusión, en nuestras condiciones de estrés térmico y sin importar el tamaño de la jaula, no se recomiendan densidades mayores a 18 conejos/m2 (34 kg/m2) al final del engorde. La productividad de la gestación y la lactancia disminuyen cuando la lactancia es mayor y se suceden varios ciclos reproductivos seguidos. Esto se debe al efecto negativo del estrés térmico sobre la vitalidad y supervivencia del embrión/gazapo. La mejora de la calidad del agua atenuó el efecto negativo del estrés térmico. Las conejas más productoras parece que son aquéllas que consiguen manejar mejor el estrés térmico.

An experimental and theoretical study of unsteady flow (gust) effects on structures

The gust wind tunnel at IDR, Universidad Politécnica de Madrid (UPM), has been enhanced and the impact of the modification has been characterized. Several flow quality configurations have been tested. The problems in measuring gusty winds with Pitot tubes have been considered. Experimental results have been obtained and compared with theoretically calculated results (based on potential flow theory). A theoretical correction term has been proposed for unsteady flow measurements obtained with Pitot tubes. The effect of unsteady flow on structures and laying bodies on the ground has been also considered. A theoretical model has been proposed for a semi-circular cylinder and experimental tests have been performed to study the unsteady flow effects, which can help in clarifying the phenomenon.

The nature of gypsum mortars for external rendering in Spain. Traditional manufacture and mineralogy

The present work focuses on gypsum mortar manufactu red in traditional kilns and used historically as exterior rendering. A documentation survey has been carried out followed by an experimental analysis using geological techniques. Conclusion shows that traditional gypsum is formed by anhydrite and inert active impurities (crystalline amorphous silica; cl ays and hydraulic phases) produced by the craft manufacture process of the system, in a kiln with a 200ºC to 1000ºC temperature interval, and continuo us fuel supply during 36 hours. Anhydrite together wit h the hydraulic phases set at consecutive time peri ods and with the presence of moisture improve the physi cal and mechanical properties of the final product. The hydration system is of great complexity and sho ws a very slow kinetics when in presence of impurities

Improving Crop Models: Incorporating New Processes, New Approaches, and Better Calibrations

Early ancestors of crop simulation models (De Wit, 1965; Monteith, 1965; Duncan et al., 1967) were born before primitive personal computers were available (e.g. Apple II released in 1977, IBM PC released in 1981). Paleo-computer programs were run in mainframes with the support of punch cards. As computers became more available and powerful, crop models evolved into sophisticated tools summarizing our understanding of how crops operate. This evolution was triggered by the need to answer new scientific questions and improve the accuracy of model simulations, especially under limiting conditions.