The effects of environmental perturbation and measurement error on estimates of the shape parameter in the theta-logistic model of population regulation

The theta-logistic is a widely used generalisation of the logistic model of regulated biological processes which is used in particular to model population regulation. Then the parameter theta gives the shape of the relationship between per-capita population growth rate and population size. Estimation of theta from population counts is however subject to bias, particularly when there are measurement errors. Here we identify factors disposing towards accurate estimation of theta by simulation of populations regulated according to the theta-logistic model. Factors investigated were measurement error, environmental perturbation and length of time series. Large measurement errors bias estimates of theta towards zero. Where estimated theta is close to zero, the estimated annual return rate may help resolve whether this is due to bias. Environmental perturbations help yield unbiased estimates of theta. Where environmental perturbations are large, estimates of theta are likely to be reliable even when measurement errors are also large. By contrast where the environment is relatively constant, unbiased estimates of theta can only be obtained if populations are counted precisely Our results have practical conclusions for the design of long-term population surveys. Estimation of the precision of population counts would be valuable, and could be achieved in practice by repeating counts in at least some years. Increasing the length of time series beyond ten or 20 years yields only small benefits. if populations are measured with appropriate accuracy, given the level of environmental perturbation, unbiased estimates can be obtained from relatively short censuses. These conclusions are optimistic for estimation of theta. (C) 2008 Elsevier B.V All rights reserved.

Factors influencing population dynamics in island and mainland populations of the Swamp Antechinus (Antechinus minimus)

Simultaneously analysing demographic processes of small mammals living in different ecological contexts may help to understand mechanisms that influence the growth and decline of these populations. The size and demography of swamp antechinus (Antechinus minimus) populations located in a coastal mainland habitat and on a small offshore island in south-eastern Australia were investigated. Large demographic differences occurred between the two ecosystems, with the island population density often 100 times greater than that on the mainland. The swamp antechinus in the mainland habitat was influenced by extrinsic climatic forces, with juvenile recruitment, individual body mass and overall population size being affected by rainfall, a factor likely to influence food availability for the species. However, the island population did not appear to be affected by drought to the same degree where allochthonous marine nutrient inputs may have offset any drought-induced reduction in primary production. Significantly greater juvenile recruitment in the island habitats combined with restricted emigration and potentially reduced predation and interspecific competition are likely to be responsible for the high population densities on the island. Although island populations appear robust, future conservation efforts should focus on mainland populations given the genetic deficiencies in the island populations.

Population genetic structure, gene flow and sex-biased dispersal in frillneck lizards (Chlamydosaurus kingii)

By using both mitochondrial and nuclear multiloci markers, we explored population genetic structure, gene flow and sex-specific dispersal of frillneck lizards (Chlamydosaurus kingii) sampled at three locations, separated by 10 to 50 km, in a homogenous savannah woodland in tropical Australia. Apart from a recombinant lizard, the mitochondrial analyses revealed two nonoverlapping haplotypes/populations, while the nuclear markers showed that the frillneck lizards represented three separate clusters/populations. Due to the small population size of the mtDNA, fixation may occur via founder effects and/or drift. We therefore suggest that either of these two processes, or a combination of the two, are the most likely causes of the discordant results obtained from the mitochondrial and the nuclear markers. In contrast to the nonoverlapping mitochondrial haplotypes, in 12 out of 74 lizards, mixed nuclear genotypes were observed, hence revealing a limited nuclear gene flow. Although gene flow should ultimately result in a blending of the populations, we propose that the distinct nuclear population structure is maintained by frequent fires resulting in local bottlenecks, and concomitant spatial separation of the frillneck lizard populations. Limited mark-recapture data and the difference in distribution of the mitochondrial and nuclear markers suggest that the mixed nuclear genotypes were caused by juvenile male-biased dispersal.

Habitat-specific demography and source-sink dynamics in a population of Siberian jays

P>1. There are a number of models describing population structure, many of which have the capacity to incorporate spatial habitat effects. One such model is the source-sink model, that describes a system where some habitats have a natality that is higher than mortality (source) and others have a mortality that exceeds natality (sink). A source can be maintained in the absence of migration, whereas a sink will go extinct. 2. However, the interaction between population dynamics and habitat quality is complex, and concerns have been raised about the validity of published empirical studies addressing source-sink dynamics. In particular, some of these studies fail to provide data on survival, a significant component in disentangling a sink from a low quality source. Moreover, failing to account for a density-dependent increase in mortality, or decrease in fecundity, can result in a territory being falsely assigned as a sink, when in fact, this density-dependent suppression only decreases the population size to a lower level, hence indicating a 'pseudo-sink'. 3. In this study, we investigate a long-term data set for key components of territory-specific demography (mortality and reproduction) and their relationship to habitat characteristics in the territorial, group-living Siberian jay (Perisoreus infaustus). We also assess territory-specific population growth rates (r), to test whether spatial population dynamics are consistent with the ideas of source-sink dynamics. 4. Although average mortality did not differ between sexes, habitat-specific mortality did. Female mortality was higher in older forests, a pattern not observed in males. Male mortality only increased with an increasing amount of open areas. Moreover, reproductive success was higher further away from human settlement, indicating a strong effect of human-associated nest predators. 5. Averaged over all years, 76% of the territories were sources. These territories generally consisted of less open areas, and were located further away from human settlement. 6. The source-sink model provides a tool for modelling demography in distinct habitat patches of different quality, which can aid in identifying key habitats within the landscape, and thus, reduce the risk of implementing unsound management decisions.

Modeling the Effects of Size on Patch Dynamics of an Inert Tracer

Mesoscale iron enrichment experiments have revealed that additional iron affects the phytoplankton productivity and carbon cycle. However, the role of initial size of fertilized patch in determining the patch evolution is poorly quantified due to the limited observational capability and complex of physical processes. Using a three-dimensional ocean circulation model, we simulated different sizes of inert tracer patches that were only regulated by physical circulation and diffusion. Model results showed that during the first few days since release of inert tracer, the calculated dilution rate was found to be a linear function with time, which was sensitive to the initial patch size with steeper slope for smaller size patch. After the initial phase of rapid decay, the relationship between dilution rate and time became an exponential function, which was also size dependent. Therefore, larger initial size patches can usually last longer and ultimately affect biogeochemical processes much stronger than smaller patches.

Interactions among chronic and acute impacts on coral recruits: the importance of size-escape thresholds

Newly settled recruits typically suffer high mortality from disturbances, but rapid growth reduces their mortality once size-escape thresholds are attained. Ocean acidification (OA) reduces the growth of recruiting benthic invertebrates, yet no direct effects on survivorship have been demonstrated. We tested whether the reduced growth of coral recruits caused by OA would increase their mortality by prolonging their vulnerability to an acute disturbance: fish herbivory on surrounding algal turf. After two months' growth in ambient or elevated CO2 levels, the linear extension and calcification of coral (Acropora millepora) recruits decreased as CO2 partial pressure (pCO2) increased. When recruits were subjected to incidental fish grazing, their mortality was inversely size dependent. However, we also found an additive effect of pCO2 such that recruit mortality was higher under elevated pCO2 irrespective of size. Compared to ambient conditions, coral recruits needed to double their size at the highest pCO2 to escape incidental grazing mortality. This general trend was observed with three groups of predators (blenny, surgeonfish, and parrotfish), although the magnitude of the fish treatment varied among species. Our study demonstrates the importance of size-escape thresholds in early recruit survival and how OA can shift these thresholds, potentially intensifying population bottlenecks in benthic invertebrate recruitment.

Individual and population-level responses to ocean acidification

Ocean acidification is predicted to have detrimental effects on many marine organisms and ecological processes. Despite growing evidence for direct impacts on specific species, few studies have simultaneously considered the effects of ocean acidification on individuals (e.g. consequences for energy budgets and resource partitioning) and population level demographic processes. Here we show that ocean acidification increases energetic demands on gastropods resulting in altered energy allocation, i.e. reduced shell size but increased body mass. When scaled up to the population level, long-term exposure to ocean acidification altered population demography, with evidence of a reduction in the proportion of females in the population and genetic signatures of increased variance in reproductive success among individuals. Such increased variance enhances levels of short-term genetic drift which is predicted to inhibit adaptation. Our study indicates that even against a background of high gene flow, ocean acidification is driving individual- and population-level changes that will impact eco-evolutionary trajectories.

Integration of socioeconomic and genetic aspects involved in the conservation of animal genetic resources = Integración de aspectos socioeconómicos y genéticos involucrados en la conservación de recursos genéticos animales

El sector ganadero está siendo gradualmente dominado por sistemas intensivos y especializados en los que los factores de producción están controlados y en los que los caracteres productivos son los criterios principales para la selección de especies y razas. Entretanto, muchos de los bienes y servicios que tradicionalmente suministraba el ganado, tales como los fertilizantes, la tracción animal o materias primas para la elaboración vestimenta y calzado están siendo reemplazados por productos industriales. Como consecuencia de ambos cambios, las razas seleccionadas intensivamente, las cuales están estrechamente ligadas a sistemas agrícolas de alta producción y altos insumos, han desplazado a muchas razas autóctonas, en las que la selección prácticamente ha cesado o es muy poco intensa. Actualmente existe una mayor conciencia social sobre la situación de las razas autóctonas y muchas funciones del ganado que previamente habían sido ignoradas están siendo reconocidas. Desde hace algunas décadas, se ha aceptado internacionalmente que las razas de ganado cumplen funciones económicas, socio-culturales, medioambientales y de seguridad alimentaria. Por ello, diferentes organismos internacionales han reconocido que la disminución de los recursos genéticos de animales domésticos (RGADs) es un problema grave y han recomendado su conservación. Aun así, la conservación de RGADs es un tema controvertido por la dificultad de valorar las funciones del ganado. Esta valoración es compleja debido que los RGADs tiene una doble naturaleza privada - pública. Como algunos economistas han subrayado, el ganado es un bien privado, sin embargo debido a algunas de sus funciones, también es un bien público. De esta forma, el aumento del conocimiento sobre valor de cada una de sus funciones facilitaría la toma de decisiones en relación a su conservación y desarrollo. Sin embargo, esta valoración es controvertida puesto que la importancia relativa de las funciones del ganado varía en función del momento, del lugar, de las especies y de las razas. El sector ganadero, debido a sus múltiples funciones, está influenciado por factores técnicos, medioambientales, sociales, culturales y políticos que están interrelacionados y que engloban a una enorme variedad de actores y procesos. Al igual que las funciones del ganado, los factores que afectan a su conservación y desarrollo están fuertemente condicionados por localización geográfica. Asimismo, estos factores pueden ser muy heterogéneos incluso dentro de una misma raza. Por otro lado, es razonable pensar que el ganadero es el actor principal de la conservación de razas locales. Actualmente, las razas locales están siendo Integration of socioeconomic and genetic aspects involved in the conservation of animal genetic resources 5 explotadas por ganaderos muy diversos bajo sistemas de producción también muy diferentes. Por todo ello, es de vital importancia comprender y evaluar el impacto que tienen las motivaciones, y el proceso de toma de decisiones de los ganaderos en la estructura genética de las razas. En esta tesis doctoral exploramos diferentes aspectos sociales, económicos y genéticos involucrados en la conservación de razas locales de ganado vacuno en Europa, como ejemplo de RGADs, esperando contribuir al entendimiento científico de este complejo tema. Nuestro objetivo es conseguir una visión global de los procesos subyacentes en la conservación y desarrollo de estas razas. Pretendemos ilustrar como se pueden utilizar métodos cuantitativos en el diseño y establecimiento de estrategias de conservación y desarrollo de RGADs objetivas y adecuadas. En primer lugar, exploramos el valor económico total (VET) del ganado analizando sus componentes públicos fuera de mercado usando como caso de estudio la raza vacuna Alistana-Sanabresa (AS). El VET de cualquier bien está formado por componentes de uso y de no-uso. Estos últimos incluyen el valor de opción, el valor de herencia y el valor de existencia. En el caso del ganado local, el valor de uso directo proviene de sus productos. Los valores de uso indirecto están relacionados con el papel que cumple las razas en el mantenimiento de los paisajes y cultura rural. El valor de opción se refiere a su futuro uso potencial y el valor de herencia al uso potencial de las generaciones venideras. Finalmente, el valor de existencia está relacionado con el bienestar que produce a la gente saber que existe un recurso específico. Nuestro objetivo fue determinar la importancia relativa que tienen los componentes fuera de mercado sobre el VET de la raza AS. Para ello evaluamos la voluntad de la gente a pagar por la conservación de la AS mediante experimentos de elección (EEs) a través de encuestas. Estos experimentos permiten valorar individualmente los distintos componentes del VET de cualquier bien. Los resultados los analizamos mediante de uso de modelos aleatorios logit. Encontramos que las funciones públicas de la raza AS tienen un valor significativo. Sus valores más importantes son el valor de uso indirecto como elemento cultural Zamorano y el valor de existencia (ambos representaron el 80% de VET). Además observamos que el valor que gente da a las funciones públicas de la razas de ganado dependen de sus características socioeconómicas. Los factores que condicionaron la voluntad a pagar para la conservación de la raza AS fueron el lugar de residencia (ciudad o pueblo), el haber visto animales de la raza o haber consumido sus productos y la actitud de los encuestados ante los conflictos entre el desarrollo económico y el medioambiente. Por otro lado, encontramos que no todo el mundo tiene una visión completa e integrada de todas las funciones públicas de la raza AS. Por este motivo, los programas o actividades de concienciación sobre su estado deberían hacer hincapié en este aspecto. La existencia de valores públicos de la raza AS implica que los ganaderos deberían recibir compensaciones económicas como pago por las funciones públicas que cumple su raza local. Las compensaciones asegurarían un tamaño de población que permitiría que la raza AS siga realizando estas funciones. Un mecanismo para ello podría ser el desarrollo del turismo rural relacionado con la raza. Esto aumentaría el valor de uso privado mientras que supondría un elemento añadido a las estrategias de conservación y desarrollo. No obstante, los ganaderos deben analizar cómo aprovechar los nichos de mercado existentes, así como mejorar la calidad de los productos de la raza prestando especial atención al etiquetado de los mismos. Una vez evaluada la importancia de las funciones públicas de las razas locales de ganado, analizamos la diversidad de factores técnicos, económicos y sociales de la producción de razas locales de ganado vacuno existente en Europa. Con este fin analizamos el caso de quince razas locales de ocho países en el contexto de un proyecto de colaboración internacional. Investigamos las diferencias entre los países para determinar los factores comunes clave que afectan a la viabilidad de las razas locales. Para ello entrevistamos mediante cuestionarios a un total de 355 ganaderos en las quince razas. Como indicador de viabilidad usamos los planes de los ganaderos de variación del tamaño de las ganaderías. Los cuestionarios incluían diferentes aspectos económicos, técnicos y sociales con potencial influencia en las dinámicas demográficas de las razas locales. Los datos recogidos los analizamos mediante distintas técnicas estadísticas multivariantes como el análisis discriminante y la regresión logística. Encontramos que los factores que afectan a la viabilidad de las razas locales en Europa son muy heterogéneos. Un resultado reseñable fue que los ganaderos de algunos países no consideran que la explotación de su raza tenga un alto valor social. Este hecho vuelve a poner de manifiesto la importancia de desarrollar programas Europeos de concienciación sobre la importancia de las funciones que cumplen las razas locales. Además los países analizados presentaron una alta variabilidad en cuanto a la importancia de los mercados locales en la distribución de los productos y en cuanto al porcentaje en propiedad del total de los pastos usados en las explotaciones. Este estudio reflejó la variabilidad de los sistemas y medios de producción (en el sentido socioeconómico, técnico y ecológico) que existe en Europa. Por ello hay que ser cautos en la implementación de las políticas comunes en los diferentes países. También encontramos que la variabilidad dentro de los países puede ser elevada debido a las diferencias entre razas, lo que implica que las políticas nacionales deber ser suficientemente flexibles para adaptarse a las peculiaridades de cada una de las razas. Por otro lado, encontramos una serie de factores comunes a la viabilidad de las razas en los distintos países; la edad de los ganaderos, la colaboración entre ellos y la apreciación social de las funciones culturales, medioambientales y sociales del ganado local. El envejecimiento de los ganaderos de razas locales no es solo un problema de falta de transferencia generacional, sino que también puede suponer una actitud más negativa hacia la inversión en las actividades ganaderas y en una menor capacidad de adaptación a los cambios del sector. La capacidad de adaptación de los ganaderos es un factor crucial en la viabilidad de las razas locales. Las estrategias y políticas de conservación comunes deben incluir las variables comunes a la viabilidad de las razas manteniendo flexibilidad suficiente para adaptarse a las especificidades nacionales. Estas estrategias y políticas deberían ir más allá de compensación económica a los ganaderos de razas locales por la menor productividad de sus razas. Las herramientas para la toma de decisiones ayudan a generar una visión amplia de la conservación y desarrollo de las razas locales. Estas herramientas abordan el diseño de estrategias de conservación y desarrollo de forma sistemática y estructurada. En la tercera parte de la tesis usamos una de estas herramientas, el análisis DAFO (Debilidades, Amenazas, Fortalezas y Oportunidades), con este propósito, reconociendo que la conservación de RGADs depende de los ganaderos. Desarrollamos un análisis DAFO cuantitativo y lo aplicamos a trece razas locales de ganado vacuno de seis países europeos en el contexto del proyecto de colaboración mencionado anteriormente. El método tiene cuatro pasos: 1) la definición del sistema; 2) la identificación y agrupación de los factores influyentes; 3) la cuantificación de la importancia de dichos factores y 4) la identificación y priorización de estrategias. Identificamos los factores utilizando multitud de agentes (multi-stakeholder appproach). Una vez determinados los factores se agruparon en una estructura de tres niveles. La importancia relativa de los cada uno de los factores para cada raza fue determinada por grupos de expertos en RGADs de los países integrados en el citado proyecto. Finalmente, desarrollamos un proceso de cuantificación para identificar y priorizar estrategias. La estructura de agrupación de factores permitió analizar el problema de la conservación desde el nivel general hasta el concreto. La unión de análisis específicos de cada una de las razas en un análisis DAFO común permitió evaluar la adecuación de las estrategias a cada caso concreto. Identificamos un total de 99 factores. El análisis reveló que mientras los factores menos importantes son muy consistentes entre razas, los factores y estrategias más relevantes son muy heterogéneos. La idoneidad de las estrategias fue mayor a medida que estas se hacían más generales. A pesar de dicha heterogeneidad, los factores influyentes y estrategias más importantes estaban ligados a aspectos positivos (fortalezas y oportunidades) lo que implica que el futuro de estas razas es prometedor. Los resultados de nuestro análisis también confirmaron la gran relevancia del valor cultural de estas razas. Las factores internos (fortalezas y debilidades) más importantes estaban relacionadas con los sistemas de producción y los ganaderos. Las oportunidades más relevantes estaban relacionadas con el desarrollo y marketing de nuevos productos mientras que las amenazas más importantes se encontraron a la hora de vender los productos actuales. Este resultado implica que sería fructífero trabajar en la motivación y colaboración entre ganaderos así como, en la mejora de sus capacidades. Concluimos que las políticas comunes europeas deberían centrarse en aspectos generales y ser los suficientemente flexibles para adaptarse a las singularidades de los países y las razas. Como ya se ha mencionado, los ganaderos juegan un papel esencial en la conservación y desarrollo de las razas autóctonas. Por ello es relevante entender que implicación puede tener la heterogeneidad de los mismos en la viabilidad de una raza. En la cuarta parte de la tesis hemos identificado tipos de ganaderos con el fin de entender cómo la relación entre la variabilidad de sus características socioeconómicas, los perfiles de las ganaderías y las dinámicas de las mismas. El análisis se ha realizado en un contexto sociológico, aplicando los conceptos de capital cultural y económico. Las tipologías se han determinado en función de factores socioeconómicos y culturales indicadores del capital cultural y capital económico de un individuo. Nuestro objetivo era estudiar si la tipología socioeconómica de los ganaderos afecta al perfil de su ganadería y a las decisiones que toman. Entrevistamos a 85 ganaderos de la raza Avileña-Negra Ibérica (ANI) y utilizamos los resultados de dichas entrevistas para ilustrar y testar el proceso. Definimos los tipos de ganaderos utilizando un análisis de clúster jerarquizado con un grupo de variables canónicas que se obtuvieron en función de cinco factores socioeconómicos: el nivel de educación del ganadero, el año en que empezó a ser ganadero de ANI, el porcentaje de los ingresos familiares que aporta la ganadería, el porcentaje de propiedad de la tierra de la explotación y la edad del ganadero. La tipología de los ganaderos de ANI resultó ser más compleja que en el pasado. Los resultados indicaron que los tipos de ganaderos variaban en muchos aspectos socioeconómicos y en los perfiles de sus Integration of socioeconomic and genetic aspects involved in the conservation of animal genetic resources 9 ganaderías. Los tipos de ganaderos determinados toman diferentes decisiones en relación a la modificación del tamaño de su ganadería y a sus objetivos de selección. Por otro lado, reaccionaron de forma diferente ante un hipotético escenario de reducción de las compensaciones económicas que les planteamos. En este estudio hemos visto que el capital cultural y el económico interactúan y hemos explicado como lo hacen en los distintos tipos de ganaderos. Por ejemplo, los ganaderos que poseían un mayor capital económico, capital cultural formal y capital cultural adquirido sobre la raza, eran los ganaderos cuyos animales tenían una mayor demanda por parte de otros ganaderos, lo cual podría responder a su mayor prestigio social dentro de la raza. Uno de los elementos claves para el futuro de la raza es si este prestigio responde a una superioridad genética de las animales. Esto ocurriría si los ganaderos utilizaran las herramientas que tienen a su disposición a la hora de seleccionar animales. Los tipos de ganaderos identificados mostraron también claras diferencias en sus formas de colaboración y en su reacción a una hipotética variación de las compensaciones económicas. Aunque algunos tipos de ganaderos mostraron un bajo nivel de dependencia a estas compensaciones, la mayoría se manifestaron altamente dependientes. Por ello cualquier cambio drástico en la política de ayudas puede comprometer el desarrollo de las razas autóctonas. La adaptación las políticas de compensaciones económicas a la heterogeneidad de los ganaderos podría aumentar la eficacia de las mismas por lo que sería interesante explorar posibilidades a este respecto. Concluimos destacando la necesidad de desarrollar políticas que tengan en cuenta la heterogeneidad de los ganaderos. Finalmente abordamos el estudio de la estructura genética de poblaciones ganaderas. Las decisiones de los ganaderos en relación a la selección de sementales y su número de descendientes configuran la estructura demográfica y genética de las razas. En la actualidad existe un interés renovado por estudiar las estructuras poblacionales debido a la influencia potencial de su estratificación sobre la predicción de valores genómicos y/o los análisis de asociación a genoma completo. Utilizamos dos métodos distintos, un algoritmo de clústeres basados en teoría de grafos (GCA) y un algoritmo de clustering bayesiano (STRUCTURE) para estudiar la estructura genética de la raza ANI. Prestamos especial atención al efecto de la presencia de parientes cercanos en la población y de la diferenciación genética entre subpoblaciones sobre el análisis de la estructura de la población. En primer lugar evaluamos el comportamiento de los dos algoritmos en poblaciones simuladas para posteriormente analizar los genotipos para 17 microsatélites de 13343 animales de 57 ganaderías distintas de raza ANI. La ANI es un ejemplo de raza con relaciones complejas. Por otro lado, utilizamos el archivo de pedigrí de la raza para estudiar el flujo de genes, calculando, entre otras cosas, la contribución de cada ganadería a la constitución genética de la raza. En el caso de las poblaciones simuladas, cuando el FST entre subpoblaciones fue suficientemente alto, ambos algoritmos, GCA y STRUCTURE, identificaron la misma estructura genética independientemente de que existieran o no relaciones familiares. Por el contrario, cuando el grado de diferenciación entre poblaciones fue bajo, el STRUCTURE identificó la estructura familiar mientras que GCA no permitió obtener ningún resultado concluyente. El GCA resultó ser un algoritmo más rápido y eficiente para de inferir la estructura genética en poblaciones con relaciones complejas. Este algoritmo también puede ser usado para reducir el número de clústeres a testar con el STRUTURE. En cuanto al análisis de la población de ANI, ambos algoritmos describieron la misma estructura, lo cual sugiere que los resultados son robustos. Se identificaron tres subpoblaciones diferenciadas que pudieran corresponderse con tres linajes distintos. Estos linajes estarían directamente relacionados con las ganaderías que han tenido una mayor contribución a la constitución genética de la raza. Por otro lado, hay un conjunto muy numeroso de individuos con una mezcla de orígenes. La información molecular describe una estructura estratificada de la población que se corresponde con la evolución demográfica de la raza. Es esencial analizar en mayor profundidad la composición de este último grupo de animales para determinar cómo afecta a la variabilidad genética de la población de ANI. SUMMARY Summary Livestock sector is gradually dominated by intensive and specialized systems where the production environment is controlled and the production traits are the main criteria for the selection of species and breeds. In the meantime, the traditional use of domestic animals for draught work, clothes and manure has been replaced by industrial products. As a consequence of both these changes, the intensively selected breeds closely linked with high-input highoutput production systems have displaced many native breeds where the selection has practically ceased or been very mild. People are now more aware of the state of endangerment among the native breeds and the previously ignored values of livestock are gaining recognition. For some decades now, the economic, socio-cultural, environmental and food security function of livestock breeds have been accepted worldwide and their loss has been recognized as a major problem. Therefore, the conservation of farm animal genetic resources (FAnGR) has been recommended. The conservation of FAnGR is controversial due to the complexity of the evaluation of its functions. This evaluation is difficult due to the nature of FAnGR both as private and public good. As some economists have highlighted, livestock animals are private goods, however, they are also public goods by their functions. Therefore, there is a need to increase the knowledge about the value of all livestock functions since to support the decision-making for the sustainable conservation and breeding of livestock. This is not straightforward since the relative importance of livestock functions depends on time, place, species and breed. Since livestock play a variety of roles, their production is driven by interrelated and everchanging economic, technical, environmental, social, cultural and political elements involving an enormous range of stakeholders. Not only FAnGR functions but also the importance of factors affecting the development and conservation of FAnGR can be very different across geographical areas. Furthermore, heterogeneity can be found even within breeds. Local breeds are nowadays raised by highly diverse farmers in equally diverse farms. It is quite reasonable to think that farmer is the major actor in the in situ conservation of livestock breeds. Thus, there is a need to understand the farmers’ motivations, decision making processes and the impact of their decisions on the genetic structure of breeds. In this PhD thesis we explore different social, economic and genetic aspects involved in the conservation of local cattle breeds, i.e. FAnGR, in Europe seeking to contribute to the scientific understanding of this complex issue. We aim to achieve a comprehensive view of the processes involved in the conservation and development of local cattle breeds and have made special efforts in discussing the implications of the research results in this respect. The final outcome of the thesis is to illustrate how quantitative methods can be exploited in designing and establishing sound strategies and programmes for the conservation and development of local livestock breeds. Firstly we explored the public non-market attributes of the total economic value (TEV) of livestock, using the Spanish Alistana-Sanabresa (AS) cattle breed as a case study. Total economic value of any good comprises both use and non-use components, where the latter include option, bequest and existence values. For livestock, the direct use values are mainly stemming from production outputs. Indirect use values relate to the role of livestock as a maintainer of rural culture and landscape. The option value is related to the potential use of livestock, the bequest values relate to the value associated with the inheritance of the resources to future generation and the existence values relate to the utility perceived by people from knowing that specific resources exist. We aimed to determine the relative importance of the non-market components of the TEV of the AS breed, the socio-economic variables that influence how people value the different components of TEV and to assess the implications of the Spanish national conservation strategy for the AS breed. To do so, we used a choice experiment (CE) approach and applied the technique to assess people’s willingness to pay (WTP) for the conservation of AS breed. The use of CE allows the valuation of the individual components of TEV for a given good. We analysed the choice data using a random parameter logit (RPL) model. AS breed was found to have a significant public good value. Its most important values were related to the indirect use value due to the maintenance of Zamorian culture and the existence value (both represent over 80% of its TEV). There were several socioeconomic variables influencing people’s valuation of the public service of the breed. In the case of AS breed, the place of living (city or rural area), having seen animals of the breed, having eaten breed products and the respondents’ attitude towards economic development – environment conflicts do influence people’s WTP for AS conservation. We also found that people do not have a complete picture of all the functions and roles that AS breed as AnGR. Therefore, the actions for increasing awareness of AS should go to that direction. The farmers will need incentives to exploit some of the public goods values and maintain the breed population size at socially desirable levels. One such mechanism could be related to the development of agritourism, which would enhance the private good value and provide an important addition to the conservation and utilisation strategy. However, the farmers need a serious evaluation on how to invest in niche product development or how to improve product quality and brand recognition. Using the understanding on the importance of the public function of local cattle we tried to depict the current diversity regarding technical, economic and social factors found in local cattle farming across Europe. To do so we focused in an international collaborative project on the case of fifteen local cattle breeds in eight European countries. We investigated the variation among the countries to detect the common key elements, which affect the viability of local breeds. We surveyed with interviews a total of 355 farms across the fifteen breeds. We used the planned herd size changes by the farmer as an indicator of breed viability. The questionnaire included several economic, technical and social aspects with potential influence on breeds’ demographic trends. We analysed the data using multivariate statistical techniques, such as discriminat analysis and logistic regression. The factors affecting a local breed’s viability were highly heterogeneous across Europe. In some countries, farmers did not recognise any high social value attached to keeping a local cattle breed. Hence there is a need to develop communication programmes across EU countries making people aware about the diversity and importance of values associated to raising local breeds. The countries were also very variable regarding the importance of local markets and the percentage of farm land owned by the farmers. Despite the country specificities, there were also common factors affecting the breed viability across Europe. The factors were from different grounds, from social, such as the age of the farmer and the social appreciation of their work, to technicalorganizational, such as the farmers’ attitude to collaborating with each other. The heterogeneity found reflects the variation in breeding systems and production environment (in the socioeconomic, technical and ecological sense) present in Europe. Therefore, caution should be taken in implementing common policies at the country level. Variability could also be rather high within countries due to breed specificities. Therefore, the national policies should be flexible to adapt to the specificities. The variables significantly associated with breed viability should be positively incorporated in the conservation strategies, and considered in developing common and/or national policies. The strategy preparation and policy planning should go beyond the provision of a general economic support to compensate farmers for the lower profitability of local breeds. Of particular interest is the observation that the opportunity for farmer collaboration and the appreciation by the society of the cultural, environmental and social role of local cattle farming were positively associated with the breed survival. In addition, farmer's high age is not only a problem of poor generation transfer but it is also a problem because it might lead to a lower attitude to investing in farming activities and to a lower ability to adapt to environment changes. The farmers’ adaptation capability may be a key point for the viability of local breeds. Decision making tools can help to get a comprehensive view on the conservation and development of local breeds. It allows us to use a systematic and structured approach for identifying and prioritizing conservation and development strategies. We used SWOT (Strengths, Weaknesses Opportunities and Threats) analysis for this purpose and recognized that many conservation and development projects rely on farmers. We developed a quantified SWOT method and applied it in the aforementioned collaborative research to a set of thirteen cattle breeds in six European countries. The method has four steps: definition of the system, identification and grouping of the driving factors, quantification of the importance of driving factors and identification and prioritization of the strategies. The factors were determined following a multi-stakeholder approach and grouped with a three level structure. FAnGR expert groups ranked the factors and a quantification process was implemented to identify and prioritize strategies. The structure of the SWOT analysis allowed analyzing the conservation problem from general down to specific perspectives. Joining breed specific analyses into a common SWOT analysis permitted comparison of breed cases across countries. We identified 99 driving factors across breeds. The across breed analysis revealed that irrelevant factors were consistent. There was high heterogeneity among the most relevant factors and strategies. The strategies increased eligibility as they lost specificity. Although the situation was very heterogeneous, the most promising factors and strategies were linked to the positive aspects (Strengths and Opportunities). Therefore, the future of the studied local breed is promising. The results of our analysis also confirmed the high relevance of the cultural value of the breeds. The most important internal factors (strengths and weaknesses) were related farmers and production systems. The most important opportunities were found in developing and marketing new products, while the most relevant threats were found in selling the current conventional products. In this regard, it should be fruitful to work on farmers’ motivation, collaboration, and capacity building. We conclude that European policies should focus on general aspects and be flexible enough to be adapted to the country and breed specificities. As mentioned, farmers have a key role in the conservation and development of a local cattle breed. Therefore, it is very relevant to understand the implications of farmer heterogeneity within a breed for its viability. In the fourth part of the thesis, we developed a general farmer typology to help analyzing the relations between farmer features and farm profiles, herd dynamics and farmers’ decision making. In the analysis we applied and used the sociological framework of economic and cultural capital and studied how the determined farmer types were linked to farm profiles and breeding decisions, among others. The typology was based on measurable socioeconomic factors indicating the economic and cultural capital of farmers. A group of 85 farmers raising the Spanish Avileña-Negra Ibérica (ANI) local cattle breed was used to illustrate and test the procedure. The farmer types were defined by a hierarchical cluster analysis with a set of canonical variables derived from the following five the socioeconomic factors: the formal educational level of the farmer, the year the farmer started keeping the ANI breed, the percentage of the total family income covered by the farm, the percentage of the total farm land owned by the farmer and the farmer’s age. The present ANI farmer types were much more complex than what they were in the past. We found that the farmer types differed in many socioeconomic aspects and in the farms profile. Furthermore, the types also differentiate farmers with respect to decisions about changing the farm size, breeding aims and stated reactions towards hypothetical subsidy variation. We have verified that economic and cultural capitals are not independent and further showed how they are interacting in the different farmer types. The farmers related to the types with high economic, institutionalized and embodied cultural capitals had a higher demand of breeding animals from others farmers of the breed, which may be related to the higher social prestige within the breed. One of the key implications of this finding for the future of the breed is whether or not the prestige of farmers is related to genetic superiority of their animals, what is to say, that it is related with a sound use of tools that farmers have available to make selection decisions. The farmer types differed in the form of collaboration and in the reactions to the hypothetical variation in subsidies. There were farmers with low dependency on subsidies, while most of them are highly dependent on subsidies. Therefore, any drastic change in the subsidy programme might have influence on the development of local breeds. The adaptation of these programme to the farmers’ heterogeneity might increase its efficacy, thus it would be interesting to explore ways of doing it. We conclude highlighting the need to have a variety of policies, which take into account the heterogeneity among the farmers. To finish we dealt with the genetic structure of livestock populations. Farmers’ decisions on the breeding animals and their progeny numbers shape the demographic and genetic structure of the breeds. Nowadays there is a renovated interest in studying the population structure since it can bias the prediction of genomic breeding values and genome wide association studies. We determined the genetic structure of ANI breed using two different methods, a graphical clustering algorithm (GCA) and a Bayesian clustering algorithm (STRUCTURE) were used. We paid particular attention to the influence that the presence of closely related individuals and the genetic differentiation of subpopulations may have on the inferences about the population structure. We first evaluated the performance of the algorithms in simulated populations. Then we inferred the genetic structure of the Spanish cattle breed ANI analysing a data set of 13343 animals (genotyped for 17 microsatellites) from 57 herds. ANI breed is an example of a population with complex relationships. We used the herdbook to study the gene flow, estimation among other things, the contribution of different herds to the genetic composition of the ANI breed. For the simulated scenarios, when FST among subpopulations was sufficiently high, both algorithms consistently inferred the correct structure regardless of the presence of related individuals. However, when the genetic differentiation among subpopulations was low, STRUCTURE identified the family based structure while GCA did not provide any consistent picture. The GCA was a fast and efficient method to infer genetic structure to determine the hidden core structure of a population with complex history and relationships. GCA could also be used to narrow down the number of clusters to be tested by STRUCTURE. Both, STRUCTURE and GCA describe a similar structure for the ANI breed suggesting that the results are robust. ANI population was found to have three genetically differentiated clusters that could correspond to three genetic lineages. These are directly related to the herds with a major contribution to the breed. In addition, ANI breed has also a large pool made of individuals with an admixture of origins. The genetic structure of ANI, assessed by molecular information, shows a stratification that corresponds to the demographic evolution of the breed. It will be of great importance to learn more about the composition of the pool and study how it is related to the existing genetic variability of the breed.

Control of time-dependent biological processes by temporally patterned input

Temporal patterning of biological variables, in the form of oscillations and rhythms on many time scales, is ubiquitous. Altering the temporal pattern of an input variable greatly affects the output of many biological processes. We develop here a conceptual framework for a quantitative understanding of such pattern dependence, focusing particularly on nonlinear, saturable, time-dependent processes that abound in biophysics, biochemistry, and physiology. We show theoretically that pattern dependence is governed by the nonlinearity of the input–output transformation as well as its time constant. As a result, only patterns on certain time scales permit the expression of pattern dependence, and processes with different time constants can respond preferentially to different patterns. This has implications for temporal coding and decoding, and allows differential control of processes through pattern. We show how pattern dependence can be quantitatively predicted using only information from steady, unpatterned input. To apply our ideas, we analyze, in an experimental example, how muscle contraction depends on the pattern of motorneuron firing.

Stochastic processes strongly influence HIV-1 evolution during suboptimal protease-inhibitor therapy

It has long been assumed that HIV-1 evolution is best described by deterministic evolutionary models because of the large population size. Recently, however, it was suggested that the effective population size (Ne) may be rather small, thereby allowing chance to influence evolution, a situation best described by a stochastic evolutionary model. To gain experimental evidence supporting one of the evolutionary models, we investigated whether the development of resistance to the protease inhibitor ritonavir affected the evolution of the env gene. Sequential serum samples from five patients treated with ritonavir were used for analysis of the protease gene and the V3 domain of the env gene. Multiple reverse transcription–PCR products were cloned, sequenced, and used to construct phylogenetic trees and to calculate the genetic variation and Ne. Genotypic resistance to ritonavir developed in all five patients, but each patient displayed a unique combination of mutations, indicating a stochastic element in the development of ritonavir resistance. Furthermore, development of resistance induced clear bottleneck effects in the env gene. The mean intrasample genetic variation, which ranged from 1.2% to 5.7% before treatment, decreased significantly (P < 0.025) during treatment. In agreement with these findings, Ne was estimated to be very small (500–15,000) compared with the total HIV-1 RNA copy number. This study combines three independent observations, strong population bottlenecking, small Ne, and selection of different combinations of protease-resistance mutations, all of which indicate that HIV-1 evolution is best described by a stochastic evolutionary model.

Gene genealogies and population variation in plants

Early in the development of plant evolutionary biology, genetic drift, fluctuations in population size, and isolation were identified as critical processes that affect the course of evolution in plant species. Attempts to assess these processes in natural populations became possible only with the development of neutral genetic markers in the 1960s. More recently, the application of historically ordered neutral molecular variation (within the conceptual framework of coalescent theory) has allowed a reevaluation of these microevolutionary processes. Gene genealogies trace the evolutionary relationships among haplotypes (alleles) with populations. Processes such as selection, fluctuation in population size, and population substructuring affect the geographical and genealogical relationships among these alleles. Therefore, examination of these genealogical data can provide insights into the evolutionary history of a species. For example, studies of Arabidopsis thaliana have suggested that this species underwent rapid expansion, with populations showing little genetic differentiation. The new discipline of phylogeography examines the distribution of allele genealogies in an explicit geographical context. Phylogeographic studies of plants have documented the recolonization of European tree species from refugia subsequent to Pleistocene glaciation, and such studies have been instructive in understanding the origin and domestication of the crop cassava. Currently, several technical limitations hinder the widespread application of a genealogical approach to plant evolutionary studies. However, as these technical issues are solved, a genealogical approach holds great promise for understanding these previously elusive processes in plant evolution.

The vesicle size of DDA:TDB liposomal adjuvants plays a role in the cell-mediated immune response but has no significant effect on antibody production

The use of cationic liposomes as experimental adjuvants for subunit peptide of protein vaccines is well documented. Recently the cationic liposome CAF01, composed of dimethyldioctadecylammonium (DDA) and trehalose dibehenate (TDB), has entered Phase I clinical trials for use in a tuberculosis (TB) vaccine. CAF01 liposomes are a heterogeneous population with a mean vesicle size of 500 nm; a strong retention of antigen at the injection site and a Th1-biassed immune response are noted. The purpose of this study was to investigate whether CAF01 liposomes of significantly different vesicle sizes exhibited altered pharmacokinetics in vivo and cellular uptake with activation in vitro. Furthermore, the immune response against the TB antigen Ag85B-ESAT-6 was followed when various sized CAF01 liposomes were used as vaccine adjuvants. The results showed no differences in vaccine (liposome or antigen) draining from the injection site, however, significant differences in the movement of liposomes to the popliteal lymph node were noted. Liposome uptake by THP-1 vitamin D3 stimulated macrophage-like cells did not show a liposome size-dependent pattern of uptake. Finally, whilst there were no significant differences in the IgG1/2 regardless of the liposome size used as a delivery vehicle for Ag85B-ESAT-6, vesicle size has a size dependent effect on cell proliferation and IL-10 production with larger liposomes (in excess of 2 µm) promoting the highest proliferation and lowest IL-10 responses, yet vesicles of ~500 nm promoting higher IFN-? cytokine production from splenocytes and higher IL-1ß at the site of injection.

A New Class of Processes for Formalizing and Generalizing Individual-Based Models: The Semi-Semi-Markov Processes

2000 Mathematics Subject Classification: 60K15, 60K20, 60G20,60J75, 60J80, 60J85, 60-08, 90B15.

Data compilation on the biological response to ocean acidification: environmental and experimental context of data sets and related literature

The exponential growth of studies on the biological response to ocean acidification over the last few decades has generated a large amount of data. To facilitate data comparison, a data compilation hosted at the data publisher PANGAEA was initiated in 2008 and is updated on a regular basis (doi:10.1594/PANGAEA.149999). By January 2015, a total of 581 data sets (over 4 000 000 data points) from 539 papers had been archived. Here we present the developments of this data compilation five years since its first description by Nisumaa et al. (2010). Most of study sites from which data archived are still in the Northern Hemisphere and the number of archived data from studies from the Southern Hemisphere and polar oceans are still relatively low. Data from 60 studies that investigated the response of a mix of organisms or natural communities were all added after 2010, indicating a welcomed shift from the study of individual organisms to communities and ecosystems. The initial imbalance of considerably more data archived on calcification and primary production than on other processes has improved. There is also a clear tendency towards more data archived from multifactorial studies after 2010. For easier and more effective access to ocean acidification data, the ocean acidification community is strongly encouraged to contribute to the data archiving effort, and help develop standard vocabularies describing the variables and define best practices for archiving ocean acidification data.

Seawater carbonate chemistry, calcification and shell size of hard clam Mercenaria mercenaria during experiments, 2011

Increasing atmospheric carbon dioxide threatens to decrease pH in the world's oceans. Coastal and estuarine calcifying organisms of significant ecological and economical importance are at risk; however, several biogeochemical processes drive pH in these habitats. In particular, coastal and estuarine sediments are frequently undersaturated with respect to calcium carbonate due to high rates of organic matter remineralization, even when overlying waters are saturated. As a result, the post-larval stages of infaunal marine bivalves must be able to deposit new shell material in conditions that are corrosive to shell. We measured calcification rates on the hard clam, Mercenaria spp.,in 5 post-larval size classes (0.39, 0.56, 0.78, 0.98, and 2.90 mm shell height) using the alkalinity anomaly method. Acidity of experimental water was controlled by bubbling with air-CO2 blends to obtain pH values of 8.02, 7.64, and 7.41, corresponding to pCO2 values of 424, 1120, and 1950 µatm. These pH values are typical of those found in many near-shore terrigenous marine sediments. Our results show that calcification rate decreased with lower pH in all 5 size classes measured. We also found a significant effect of size on calcification rate, with the smaller post-larval sizes unable to overcome dissolution pressure. Increased calcification rate with size allowed the larger sizes to overcome dissolution pressure and deposit new shell material under corrosive conditions. Size dependency of pH effects on calcification is likely due to organogenesis and developmental shifts in shell mineralogy occurring through the post-larval stage. Furthermore, we found significantly different calcification rates between the 2 sources of hard clams we used for these experiments, most likely due to genotypic differences. Our findings confirm the susceptibility of the early life stages of this important bivalve to decreasing pH and reveal mechanisms behind the increased mortality in post-larval juvenile hard clams related to dissolution pressure, that has been found in previous studies.