Regulación de la hidratación y la turgencia foliares por mecanismos evitadores del estrés, y resistencia a déficit hídrico en vid

El transporte del agua en las plantas es impulsado por diferencias de energía libre entre el suelo y la atmósfera, y está regulado por mecanismos biológicos evitadores, como el cierre estomático. La hidratación y la turgencia foliares resultan del equilibrio entre ΨL del apoplasto, el potencial osmótico del simplasto y la elasticidad de los tejidos. Sobre esta base se conjeturó que las interacciones de los mecanismos evitadores del estrés hídrico de la planta tienen un rol clave en la definición de su resistencia a déficit hídrico. Para probar esta hipótesis se construyó un modelo mecanístico basado en las leyes del flujo de savia de Van de Honert, de difusión de Fick, de elasticidad de Hooke, la ecuación de Gardner para el flujo del agua en la rizósfera y el modelo de conductancia estomática (gs) de Buckley. Mediante el modelo se demostró teóricamente que la hidratación y la turgencia foliares dependen de la oferta de agua edáfica (representada por el potencial hídrico del suelo) y de la demanda evaporativa de la atmósfera (representada por la radiación absorbida, la temperatura del aire, la velocidad del viento y el déficit de presión de vapor de la atmósfera). También que los mecanismos evitadores del estrés hídrico -i.e., conductancia hidráulica de la planta, conductancia estomática, elasticidad del tejido y potencial osmótico a turgencia máxima- son todos necesarios para determinar la hidratación y la turgencia foliares. El modelo también demostró que la conductancia hidráulica suelo-hoja (kL) depende de la fracción de agua edáfica transpirable (FTSW) con un patrón de decaimiento sigmoide, a medida que el suelo se seca. Esto implica que las variables que dependen en parte de kL (i.e., gs, transpiración, fotosíntesis y superficie foliar) también dependen de FTSW con el mismo patrón. El modelo se probó experimentalmente a distintos niveles de humedad edáfica (desde déficit hídrico nulo, hasta severo) en cinco variedades de vid y mostró un poder predictivo superior al 90%. En todas las variedades las gs se asociaron linealmente con las kL observadas, al considerar todas las situaciones de déficit hídrico en conjunto, si bien la pendiente de estas relaciones fueron distintas en cada variedad. La contrastación experimental mostró que, en una escala de tiempo de varios meses, las variedades más evitadoras -i.e., Grenache y Cereza- mantuvieron mayor kL, ajuste osmótico y rigidez de los tejidos y una menor pendiente de la relación de gs vs. kL, que las variedades menos evitadoras -i.e., Malbec y Syrah-. La menor pendiente de la relación entre gs y kL, en las variedades más evitadoras, estuvo asociada a una mayor cantidad de estomas, en relación con la cantidad de células epidérmicas. Los variedades más evitadoras bajo déficit hídrico moderado -i.e., con una fracción de agua edáfica transpirable entre 0,6 y 0,4- tuvieron mayor superficie foliar y produjeron más biomasa, favoreciendo raíces profundas y densas, y ahorrando agua. Chardonnay mantuvo una alta hidratación y turgencia a expensas de un alto gasto de agua debido a que privilegiaba una alta kL por sobre el ajuste estomático, por lo que no podría considerarse en forma estricta como muy evitadora.

Influencia de seis portainjertos de vid sobre el comportamiento vitícola de la cv. Malbec y estudio de las relaciones hídricas que se establecen

El uso de portainjertos en la vid se ha difundido por su resistencia a filoxera y nemátodos, pero también por su tolerancia a condiciones adversas del suelo. Por otro lado, los portainjertos modifican las relaciones fuente-destino, influyendo en el comportamiento vegetativo y reproductivo de las plantas y en la composición de la uva, lo cuál puede ser utilizado como una herramienta de manejo agronómico. A fin de evaluar si existe un comportamiento diferencial de los portainjertos en cuanto a expresión vegetativa, vigor, rendimiento y composición de la uva, y explicar dichas diferencias en términos de exploración radical, relaciones hídricas, asimilación de carbono, eficiencia en el uso del agua y partición de asimilados se realizó un ensa-yo a campo de cv. Malbec sobre seis portainjertos (3309 C, 1103 P, 140 Ru, SO4, Harmony y Cereza) y a pie franco. Los portainjertos 140 Ru, 1103 P y SO4 tuvieron una mayor tendencia a la producción de uva (mayor Índice de Ravaz), y Franco, Cereza y 3309 C a vegetar, mostrando Harmony una situación intermedia. Las ba-yas sobre el pie Cereza tuvieron un mayor peso (1,96 g) que sobre Harmony (1,75 g). No se encontraron diferencias en los polifenoles de las bayas entre portainjertos. La fotosíntesis de la planta entera (Amax) de Franco, 1103 P y SO4 fue mayor que la de Harmony. La conductancia hidráulica foliar específica (kL) de Harmony fue me-nor que la de Cereza, y su conductancia hidráulica (kH) fue menor que la de Franco, Cereza y SO4. El número de raíces totales de 140 Ru fue mayor que el de 1103 P, SO4 y Harmony. El portainjerto 140 Ru se destacó por privilegiar el desarrollo radi-cal y reproductivo sobre el vegetativo, y por su mayor eficiencia en el uso del agua (EUA). Las diferencias entre portainjertos pueden ser explicadas en parte por dife-rencias en la kL que a su vez incide en el estado hídrico de las plantas (ΨL). De ma-nera que cuando la kL es más baja, el ΨL es menor (i.e., Harmony), y cuando la kL es más alta, el ΨL es mayor (i.e., Franco y Cereza). Mayores ΨL se asocian con mayores superficies foliares.

An example of integration between en economic and an hydrologic model in the framework of water resource management problems

Growing scarcity, increasing demand and bad management of water resources are causing weighty competition for water and consequently managers are facing more and more pressure in an attempt to satisfy users? requirement. In many regions agriculture is one of the most important users at river basin scale since it concentrates high volumes of water consumption during relatively short periods (irrigation season), with a significant economic, social and environmental impact. The interdisciplinary characteristics of related water resources problems require, as established in the Water Framework Directive 2000/60/EC, an integrated and participative approach to water management and assigns an essential role to economic analysis as a decision support tool. For this reason, a methodology is developed to analyse the economic and environmental implications of water resource management under different scenarios, with a focus on the agricultural sector. This research integrates both economic and hydrologic components in modelling, defining scenarios of water resource management with the goal of preventing critical situations, such as droughts. The model follows the Positive Mathematical Programming (PMP) approach, an innovative methodology successfully used for agricultural policy analysis in the last decade and also applied in several analyses regarding water use in agriculture. This approach has, among others, the very important capability of perfectly calibrating the baseline scenario using a very limited database. However one important disadvantage is its limited capacity to simulate activities non-observed during the reference period but which could be adopted if the scenario changed. To overcome this problem the classical methodology is extended in order to simulate a more realistic farmers? response to new agricultural policies or modified water availability. In this way an economic model has been developed to reproduce the farmers? behaviour within two irrigation districts in the Tiber High Valley. This economic model is then integrated with SIMBAT, an hydrologic model developed for the Tiber basin which allows to simulate the balance between the water volumes available at the Montedoglio dam and the water volumes required by the various irrigation users.

Effect of row orientation on soil water content and vine water status on a Cabernet franc vineyard in Madrid, spain

The aim of this study was to evaluate the effects of row orien¬tation on vine and soil water status in an irrigated vineyard. The trial was developed during 2006, 2007 and 2008, in the South East region of Madrid (Spain) on 5-year old Cabernet franc grapevines (Vitis vinifera L.) grafted onto 140Ru. Plant spacing was 2.5 m x 1.5 m and vines were trained to a VSP. Four orientations were stu¬died: North-South (N-S), East-West (E-W), Northeast-Southwest (N+45) and North-South +20o (N+20). Irrigation (0.4•ET0) started when shoot growth stopped. Soil water availability was measured using a TDR technique with forty buried probes. Row orientation did not have any effect on water consumption in the vineyard. At maturity, leaf water potential was measured at predawn, early mor¬ning, midday and 14:00 solar time, on both canopy sides - sun and shade – ; the early morning measurement was the one that better differentiated treatments. Leaf water potential was a good indica¬tor of plant water status. Differences between (N-S and E-W) and (N+20 and N+45) treatments were obtained both on sun and shade canopy sides, N+20 and N+45 having lower leaf water potentials then drier leaves. The water stress integral shows that N-S and E-W reach the end of maturation with a greater level of hydration than N+45 and N+20. As a whole, N+45 and N+20 orientations, without affecting too much the soil available water content, induce regularly more water stress to the vine at some periods, probably due to an higher sunlight interception in early morning which makes water limitation for the vine more early and thus more severe during the day.

Latin American Agricultural Trade: The Role of the WTO in Sustainable Virtual Water Flows, European Association of Agricultural Economists

International agricultural trade has been growing significantly during the last decade. Many countries rely on imports to ensure adequate food supplies to the people. A few are becoming food baskets of the world. This process raises issues about the food security in depending countries and potentially unsustainable land and water use in exporting countries. In this paper, we analyse the impacts of amplified farm trade on natural resources, especially water. Farm exports and imports of five Latin America countries (Brazil, Argentina, Mexico, Peru and Chile) are examined carefully. A preliminary analysis indicates that virtual water imports can save valuable water resources in water-short countries, such as Mexico and Chile. Major exporting countries, including Brazil and Argentina, have become big exporters due to abundant natural resource endowments. The opportunity costs of agricultural production in those countries are identified as being low, because of the predominant green water use. It is concluded that virtual water trade can be a powerful tool to alleviate water stress in semi-arid countries. However, for exporting nations a sustainable water use can only be guaranteed if environmental production costs are fully reflected in the commodity prices. There is no basis for erecting environmental trade tariffs on exporters though. Setting up legal foundations for them in full compliance with WTOs processes would be a daunting task.

Evaluation of Spain's Water-Energy Nexus

This paper explores the water-energy nexus of Spain and offers calculations for both the energy used in the water sector and the water required to run the energy sector. The article takes a prospective approach, offering evaluations of policy objectives for biofuels and expected renewable energy sources. Approximately 5.8% of total electricity demand in Spain is due to the water sector. Irrigated agriculture is one of the Spanish water sectors that show the largest growth in energy requirements. Searches for more efficient modes of farm water use, urban waste water treatment, and the use of desalinated water must henceforth include the energy component. Furthermore, biofuel production, to the levels targeted for 2020, would have an unbearable impact on the already stressed water resources in Spain. However, growing usage of renewable energy sources is not threatened by water scarcity, but legislative measures in water allocation and water markets will be required to meet the requirements of using these sources. Some of these measures, which are pushed by regional governments, are discussed in concluding sections.

River Restoration in Spain: Theoretical and Practical Approach in the Context of the European Water Framework Directive.

River restoration is becoming a priority in many countries because of increasing the awareness of environmental degradation. In Europe, the EU Water Framework Directive (WFD) has significantly reinforced river restoration, encouraging the improvement of ecological status for water bodies. To fulfill the WFD requirements, the Spanish Ministry of the Environment developed in 2006 a National Strategy for River Restoration whose design and implementation are described in this paper. At the same time many restoration projects have been conducted, and sixty of them have been evaluated in terms of stated objectives and pressures and implemented restoration measures. Riparian vegetation enhancement, weir removal and fish passes were the most frequently implemented restoration measures, although the greatest pressures came from hydrologic alteration caused by flow regulation for irrigation purposes. Water deficits in quantity and quality associated with uncontrolled water demands seriously affect Mediterranean rivers and represent the main constraint to achieving good ecological status of Spanish rivers, most of them intensively regulated. Proper environmental allocation of in-stream flows would need deep restrictions in agricultural water use which seem to be of very difficult social acceptance. This situation highlights the need to integrate land-use and rural development policies with water resources and river management, and identifies additional difficulties in achieving the WFD objectives and good ecological status of rivers in Mediterranean countries.

Electrical Power Fluctuations in a Network of DC/AC inverters in a Large PV Plant: relationship between correlation, distance and time scale

This paper analyzes the correlation between the fluctuations of the electrical power generated by the ensemble of 70 DC/AC inverters from a 45.6 MW PV plant. The use of real electrical power time series from a large collection of photovoltaic inverters of a same plant is an impor- tant contribution in the context of models built upon simplified assumptions to overcome the absence of such data. This data set is divided into three different fluctuation categories with a clustering proce- dure which performs correctly with the clearness index and the wavelet variances. Afterwards, the time dependent correlation between the electrical power time series of the inverters is esti- mated with the wavelet transform. The wavelet correlation depends on the distance between the inverters, the wavelet time scales and the daily fluctuation level. Correlation values for time scales below one minute are low without dependence on the daily fluctuation level. For time scales above 20 minutes, positive high correlation values are obtained, and the decay rate with the distance depends on the daily fluctuation level. At intermediate time scales the correlation depends strongly on the daily fluctuation level. The proposed methods have been implemented using free software. Source code is available as supplementary material.

Exploring the feasibility of integrating water issues into the CAPRI model

Although numerous modelling efforts have integrated food and water considerations at the farm or river basin level, very few agro-economic models are able to jointly assess water and food policies at the global level. The present report explores the feasibility of integrating water considerations into the CAPRI model. First, a literature review of modelling approaches integrating food and water issues has been conducted. Three agro-economic models, IMPACT, WATERSIM and GLOBIOM, have been analysed in detail. In addition, biophysical and hydrological models estimating agricultural water use have also been studied, in particular the global hydrological model WATERGAP and the LISFLOOD model. Thanks to the programming approach of its supply module, CAPRI shows a high potentiality to integrate environmental indicators as well as to enter new resource constraints (land potentially irrigated, irrigation water) and input-output relationships. At least in theory, the activity-based approach of the regional programming model in CAPRI allows differentiating between rainfed and irrigated activities. The suggested approach to include water into the CAPRI model involves creating an irrigation module and a water use module. The development of the CAPRI water module will enable to provide scientific assessment on agricultural water use within the EU and to analyze agricultural pressures on water resources. The feasibility of the approach has been tested in a pilot case study including two NUTS 2 regions (Andalucia in Spain and Midi-Pyrenees in France). Preliminary results are presented, highlighting the interrelations between water and agricultural developments in Europe. As a next step, it is foreseen to further develop the CAPRI water module to account for competition between agricultural and non-agricultural water use. This will imply building a water use sub-module to compute water use balances.

The Spanish water ?pressure cooker?: Threading the interplay between resource resilient water governance outcomes by strengthening the robustness of water governance processes.

This paper uses the metaphor of a pressure cooker to highlight how water problems in Spain are highly geographical and sectorial in nature, with some specific hotspots which raise the temperature of the whole water complex system, turning many potentially solvable water problems into ?wicked problems?. The paper discusses the tendency for water governance to be hydrocentric, when often the drivers and in turn the ?solutions? to Spanish water problems lie outside the water sphere. The paper analyzes of the current water governance system by looking at water governance as both a process, and its key attributes like participation, trans- parency, equity and rule of law, as well as an analysis of water governance as an outcome by looking at efficiency and sustainability of water use in Spain. It concludes on the need to have a deeper knowledge on the interactions of water governance as a process and as an outcome and potential synergies and arguing that water governance is an inherently political process which calls for strengthening the capacity of the system by looking at the interactions of these different governance attributes.

Participatory modelling to support decision making in water management under uncertainty: Two comparative case studies in the Guadiana river basin, Spain.

A participatory modelling process has been conducted in two areas of the Guadiana river (the upper and the middle sub-basins), in Spain, with the aim of providing support for decision making in the water management field. The area has a semi-arid climate where irrigated agriculture plays a key role in the economic development of the region and accounts for around 90% of water use. Following the guidelines of the European Water Framework Directive, we promote stakeholder involvement in water management with the aim to achieve an improved understanding of the water system and to encourage the exchange of knowledge and views between stakeholders in order to help building a shared vision of the system. At the same time, the resulting models, which integrate the different sectors and views, provide some insight of the impacts that different management options and possible future scenarios could have. The methodology is based on a Bayesian network combined with an economic model and, in the middle Guadiana sub-basin, with a crop model. The resulting integrated modelling framework is used to simulate possible water policy, market and climate scenarios to find out the impacts of those scenarios on farm income and on the environment. At the end of the modelling process, an evaluation questionnaire was filled by participants in both sub-basins. Results show that this type of processes are found very helpful by stakeholders to improve the system understanding, to understand each others views and to reduce conflict when it exists. In addition, they found the model an extremely useful tool to support management. The graphical interface, the quantitative output and the explicit representation of uncertainty helped stakeholders to better understand the implications of the scenario tested. Finally, the combination of different types of models was also found very useful, as it allowed exploring in detail specific aspects of the water management problems.

Thermographic imaging: assessment of drought and heat tolerance in Spanish germplasm of Brachypodium distachyon.

The annual grass Brachypodium distachyon has been recently recognized as the model plant for functional genomics of temperate grasses, including cereals of economic relevance like wheat and barley. Sixty-two lines of B. distachyon were assessed for response to drought stress and heat tolerance. All these lines, except the reference genotype BD21, derive from specimens collected in 32 distinct locations of the Iberian Peninsula, covering a wide range of geo- climatic conditions. Sixteen lines of Brachypodium hybridum, an allotetraploid closely related to B. distachyon were used as reference of abiotic-stress well-adapted genotypes. Drought tolerance was assessed in a green-house trial. At the rosette-stage, no irrigation was applied to treated plants whereas their replicates at the control were maintained well watered during all the experiment. Thermographic images of treated and control plants were taken after 2 and 3 weeks of drought treatment, when stressed plants showed medium and extreme wilting symptoms. The mean leaf temperature of stressed (LTs) and control (LTc) plants was estimated based upon thermographic records from selected pixels (183 per image) that strictly correspond to leaf tissue. The response to drought was based on the analysis of two parameters: LTs and the thermal difference (TD) between stressed and control plants (LTs – LTc). The response to heat stress was based on LTc. Comparison of the mean values of these parameters showed that: 1) Genotypes better adapted to drought (B. hybridum lines) presented a higher LTs and TD than B. distachyon lines. 2) Under high temperature conditions, watered plants of B. hybridum lines maintained lower LTc than those of B. distachyon. Those results suggest that in these species adaptation to drought is linked to a more efficient stomata regulation: under water stress stomata are closed, increasing foliar temperature but also water use efficiency by reducing transpiration. With high temperature and water availability the results are less definite, but still seems that opening stomata allow plants to increase transpiration and therefore to diminish foliar temperature.

Water option contracts for reducing water supply risks: an application to the Tagus-Segura Transfer

El agua es un recurso cada vez más escaso y valioso. Por ello, los recursos hídricos disponibles deben asignarse de una forma eficiente entre los diferentes usos. El cambio climático aumentará la frecuencia y severidad de los eventos extremos, y podría incrementar la demanda de agua de los cultivos. El empleo de mecanismos flexibles de asignación de agua puede ser imprescindible para hacer frente a este aumento en la variabilidad del balance hídrico y para asegurar que los riesgos de suministro, y no solo los recursos, son compartidos de manera eficiente entre los usuarios. Los mercados de agua permiten la reasignación de los recursos hídricos, favoreciendo su transferencia desde los usos de menor a los de mayor valor. Diferentes tipos de mercados de agua se han establecido en diferentes partes del mundo, ayudando a los participantes a afrontar los problemas de escasez de agua en esas zonas. En España, los intercambios de agua están permitidos desde 1999, aunque la participación de los usuarios en el mercado ha sido limitada. Hay varios aspectos de los mercados de agua en España que deben mejorarse. Esta tesis, además de proponer una serie de cambios en el marco regulatorio, propone la introducción de contratos de opción de agua como una posible mejora. La principal ventaja de este tipo de contratos es la estabilidad legal e institucional que éstos proporcionan tanto a compradores como vendedores. Para apoyar esta propuesta, se han llevado a cabo diferentes análisis que muestran el potencial de los contratos de opción como herramienta de reducción del riesgo asociado a una oferta de agua inestable. La Cuenca del Segura (Sureste de España), la Cuenca del Tajo y el Acueducto Tajo- Segura han sido seleccionados como casos de estudio. Tres análisis distintos aplicados a dicha región se presentan en esta tesis: a) una evaluación de los contratos de opción como mecanismo para reducir los riesgos de disponibilidad de agua sufridos por los regantes en la Cuenca del Segura; b) un marco teórico para analizar las preferencias de los regantes por diferentes mecanismos de gestión del riesgo de disponibilidad de agua, su disposición a pagar por ellos y los precios aproximados de estos instrumentos (seguro de sequía y contratos de opción de agua); y c) una evaluación del papel de los contratos de opción en las decisiones de aprovisionamiento de agua de una comunidad de regantes ante una oferta de agua incierta. Los resultados muestran el potencial de reducción del riesgo de los contratos de opción para regantes en España, pero pueden ser extrapolados a otros sectores o regiones. Las principales conclusiones de esta tesis son: a) la agricultura será uno de los sectores más afectados por el cambio climático. Si los precios del agua aumentan, la rentabilidad de los cultivos puede caer hasta niveles negativos, lo que podría dar lugar al abandono de cultivos de regadío en algunas zonas de España. Las políticas de cambio climático y de agua deben estar estrechamente coordinadas para asegurar un uso de agua eficiente y la rentabilidad de la agricultura; b) aunque los mercados de agua han ayudado a algunos usuarios a afrontar problemas de disponibilidad del recurso en momentos de escasez, hay varios aspectos que deben mejorarse; c) es necesario desarrollar mercados de agua más flexibles y estables para garantizar una asignación eficiente de los recursos entre los usuarios de agua; d) los resultados muestran los beneficios derivados del establecimiento de un contrato de opción entre usuarios de agua del Tajo y del Segura para reducir el riesgo de disponibilidad de agua en la cuenca receptora; e) la disposición a pagar de los regantes por un contrato de opción de agua o un seguro de sequía hidrológica, que representa el valor que tienen estos mecanismos para aquellos usuarios de agua que se enfrentan a riesgos relacionados con la disponibilidad del recurso, es consistente con los resultados obtenidos en estudios previos y superior al precio de mercado de estos instrumentos, lo que favorece la viabilidad de estos mecanismos de gestión del riesgo ; y f) los contratos de opción podrían ayudar a optimizar las decisiones de aprovisionamiento de agua bajo incertidumbre, proporcionando más estabilidad y flexibilidad que los mercados temporales de agua. ABSTRACT Water is becoming increasingly scarce and valuable. Thus, existing water resources need to be efficiently allocated among users. Climate change is expected to increase the frequency and severity of extreme events, and it may also increase irrigated crops' water demand. The implementation of flexible allocation mechanisms could be essential to cope with this increased variability of the water balance and ensure that supply risks, and not only water resources, are also efficiently shared and managed. Water markets allow for the reallocation of water resources from low to high value uses. Different water trading mechanisms have been created in different parts of the world and have helped users to alleviate water scarcity problems in those areas. In Spain, water trading is allowed since 1999, although market activity has been limited. There are several issues in the Spanish water market that should be improved. This thesis, besides proposing several changes in the legislative framework, proposes the introduction of water option contracts as a potential improvement. The main advantage for both buyer and seller derived from an option contract is the institutional and legal stability it provides. To support this proposal, different analyses have been carried out that show the potential of option contracts as a risk reduction tool to manage water supply instability. The Segura Basin (Southeast Spain), the Tagus Basin and the Tagus-Segura inter-basin Transfer have been selected as the case study. Three different analyses applied to this region are presented in this thesis: a) an evaluation of option contracts as a mechanisms to reduce water supply availability risks in the Segura Basin; b) a theoretical framework for analyzing farmer’s preferences for different water supply risk management tools and farmers’ willingness to pay for them, together with the assessment of the prices of these mechanisms (drought insurance and water option contracts); and c) an evaluation of the role of option contracts in water procurement decisions under uncertainty. Results show the risk-reduction potential of option contracts for the agricultural sector in Spain, but these results can be extrapolated to other sectors or regions. The main conclusions of the thesis are: a) agriculture would be one of the most affected sectors by climate change. With higher water tariffs, crop’s profitability can drop to negative levels, which may result in the abandoning of the crop in many areas. Climate change and water policies must be closely coordinated to ensure efficient water use and crops’ profitability; b) although Spanish water markets have alleviated water availability problems for some users during water scarcity periods, there are several issues that should be improved; c) more flexible and stable water market mechanisms are needed to allocate water resources and water supply risks among competing users; d) results show the benefits derived from the establishment of an inter-basin option contract between water users in the Tagus and the Segura basins for reducing water supply availability risks in the recipient area; e) irrigators’ willingness to pay for option contracts or drought insurance, that represent the value that this kind of trading mechanisms has for water users facing water supply reliability problems, are consistent with results obtained in previous works and higher than the prices of this risk management tools, which shows the feasibility of these mechanisms; and f) option contracts would help to optimize water procurement decisions under uncertainty, providing more flexibility and stability than the spot market.

Habitat management for conservation of pollinators in agro-ecosystems of Central Spain

La gestión de hábitat orientada a la conservación de polinizadores en los agro-ecosistemas requiere una selección de especies vegetales atendiendo fundamentalmente a dos criterios: i) el potencial atractivo de sus flores a los polinizadores; y ii) la simplicidad en su manejo agronómico. Además de estas premisas, es necesario considerar la capacidad invasora de estas especies vegetales, debido a que algunas de las más atractivas pueden resultar invasoras en determinados agro-ecosistemas. Por lo tanto, es preciso determinar qué especies vegetales son las más indicadas para ser implementadas en cada agro-ecosistema. En la presente tesis doctoral se plantea la búsqueda de las especies vegetales adecuadas para atraer polinizadores en los agro-ecosistemas del centro de España. En una primera aproximación, se ha evaluado la atracción y expansión espacial (potencial invasivo) de seis plantas perennes de la familia Lamiaceae (aromáticas), elegidas por ser nativas de la región mediterránea. La elección de las especies vegetales se ha llevado a cabo con el fin de crear márgenes funcionales basados en la mezcla de especies vegetales con distintos periodos de floración, de modo que prolonguen la disponibilidad de recursos florales en el tiempo. Tras un primer año dedicado al establecimiento de las especies aromáticas, en los dos años siguientes se ha estudiado la atracción individual y combinada de las especies vegetales sobre los polinizadores, y como ésta se ve afectada por la densidad y la morfología floral, utilizando para ello un diseño experimental en bloques al azar. Los resultados de este estudio han puesto de manifiesto que la morfología floral no tuvo influencia sobre la atracción de las especies vegetales, pero si la densidad floral, puesto que las especies vegetales con mayor densidad de flores (Nepeta tuberosa e Hyssopus officinalis) han mostrado mayor atracción a polinizadores. Cabe destacar que de las seis especies consideradas, dos especies de verano (Melissa officinalis y Thymbra capitata) no han contribuido de forma efectiva a la atracción de la mezcla hacia los polinizadores, mostrando una reducción significativa de este parámetro respecto a las otras especies aromáticas a lo largo del verano. Se ha observado que ninguna de las especies aromáticas evaluadas ha mostrado tendencia invasora a lo largo del estudio. En base a estos resultados, se puede concluir que entre las especies aromáticas estudiadas, N. tuberosa, H. officinalis y Salvia verbenaca son las que ofrecen mayor potencial para ser utilizadas en la conservación de polinizadores. De forma similar al caso de las plantas aromáticas, se ha llevado a cabo una segunda experimentación que incluía doce plantas anuales con floración de primavera, en la que se evaluó la atracción a polinizadores y su comportamiento agronómico. Este estudio con especies herbáceas se ha prolongado durante dos años, utilizando un diseño experimental de bloques aleatorios. Las variables analizadas fueron: el atractivo de las distintas especies vegetales a los polinizadores, su eficiencia de atracción (calculada como una combinación de la duración de la floración y las visitas de insectos), su respuesta a dos tipos de manejo agronómico (cultivo en mezcla frente a monocultivo; laboreo frente a no-laboreo) y su potencial invasivo. Los resultados de esta segunda experimentación han mostrado que las flores de Borago officinalis, Echium plantagineum, Phacelia tanacetifolia y Diplotaxis tenuifolia son atractivas a las abejas, mientras que las flores de Calendula arvensis, Coriandrum sativum, D. tenuifolia y Lobularia maritima son atractivas a los sírfidos. Con independencia del tipo de polinizadores atraídos por cada especie vegetal, se ha observado una mayor eficiencia de atracción en parcelas con monocultivo de D. tenuifolia respecto a las parcelas donde se cultivó una mezcla de especies herbáceas, si bien en estas últimas se observó mayor eficiencia de atracción que en la mayoría de parcelas mono-específicas. Respecto al potencial invasivo de las especies herbáceas, a pesar de que algunas de las más atractivas a polinizadores (P. tanacetifolia and C. arvensis) mostraron tendencia a un comportamiento invasor, su capacidad de auto-reproducción se vio reducida con el laboreo. En resumen, D. tenuifolia es la única especie que presentó una alta eficiencia de atracción a distintos tipos de polinizadores, conjuntamente con una alta capacidad de auto-reproducción pero sin mostrar carácter invasor. Comparando el atractivo de las especies vegetales utilizadas en este estudio sobre los polinizadores, D. tenuifolia es la especie más recomendable para su cultivo orientado a la atracción de polinizadores en agro-ecosistemas en el centro de España. Esta especie herbácea, conocida como rúcula, tiene la ventaja añadida de ser una especie comercializada para el consumo humano. Además de su atractivo a polinizadores, deben considerarse otros aspectos relacionados con la fisiología y el comportamiento de esta especie vegetal en los agro-ecosistemas antes de recomendar su cultivo. Dado que el cultivo en un campo agrícola de una nueva especie vegetal implica unos costes de producción, por ejemplo debidos a la utilización de agua de riego, es necesario evaluar el incremento en dichos costes en función de demanda hídrica específica de esa especie vegetal. Esta variable es especialmente importante en zonas dónde se presentan sequías recurrentes como es el caso del centro y sur-este de la península Ibérica. Este razonamiento ha motivado un estudio sobre los efectos del estrés hídrico por sequía y el estrés por déficit moderado y severo de riego sobre el crecimiento y floración de la especie D. tenuifolia, así como sobre la atracción a polinizadores. Los resultados muestran que tanto el crecimiento y floración de D. tenuifolia como su atracción a polinizadores no se ven afectados si la falta de riego se produce durante un máximo de 4 días. Sin embargo, si la falta de riego se extiende a lo largo de 8 días o más, se observa una reducción significativa en el crecimiento vegetativo, el número de flores abiertas, el área total y el diámetro de dichas flores, así como en el diámetro y longitud del tubo de la corola. Por otro lado, el estudio pone de manifiesto que un déficit hídrico regulado permite una gestión eficiente del agua, la cual, dependiendo del objetivo final del cultivo de D. tenuifolia (para consumo o solo para atracción de polinizadores), puede reducir su consumo entre un 40 y un 70% sin afectar al crecimiento vegetativo y desarrollo floral, y sin reducir significativamente el atractivo a los polinizadores. Finalmente, esta tesis aborda un estudio para determinar cómo afecta el manejo de hábitat a la producción de los cultivos. En concreto, se ha planteado una experimentación que incluye márgenes mono-específicos y márgenes con una mezcla de especies atractivas a polinizadores, con el fin de determinar su efecto sobre la producción del cultivo de cilantro (C. sativum). La elección del cultivo de cilantro se debe a que requiere la polinización de insectos para su reproducción (aunque, en menor medida, puede polinizarse también por el viento), además de la facilidad para estimar su producción en condiciones semi-controladas de campo. El diseño experimental consistía en la siembra de márgenes mono-específicos de D. tenuifolia y márgenes con mezcla de seis especies anuales situados junto al cultivo de cilantro. Estos cultivos con márgenes florales fueron comparados con controles sin margen floral. Además, un segundo grupo de plantas de cilantro situadas junto a todos los tratamientos, cuyas flores fueron cubiertas para evitar su polinización, sirvió como control para evaluar la influencia de los polinizadores en la producción del cultivo. Los resultados muestran que la presencia de cualquiera de los dos tipos de margen floral mejora el peso y el porcentaje de germinación de las semillas de cilantro frente al control sin margen. Si se comparan los dos tipos de margen, se ha observado un mayor número de semillas de cilantro junto al margen con mezcla de especies florales respecto al margen mono-específico, probablemente debido al mayor número visitas de polinizadores. Puesto que el experimento se realizó en condiciones de campo semi-controladas, esto sugiere que las visitas de polinizadores fueron el factor determinante en los resultados. Por otro lado, los resultados apuntan a que la presencia de un margen floral (ya sea mono-especifico o de mezcla) en cultivos de pequeña escala puede aumentar la producción de cilantro en más de un 200%, al tiempo que contribuyen a la conservación de los polinizadores. ABSTRACT Habitat management, aimed to conserve pollinators in agro-ecosystems, requires selection of the most suitable plant species in terms of their attractiveness to pollinators and simplicity of agronomic management. However, since all flowers are not equally attractive to pollinators and many plant species can be weedy or invasive in the particular habitat, it is important to test which plant species are the most appropriate to be implemented in specific agro-ecosystems. For that reason, this PhD dissertation has been focused on determination of the most appropriate aromatic and herbaceous plants for conservation of pollinators in agro-ecosystems of Central Spain. Therefore, in a first approximation, spatial expansion (i.e. potential weediness) and attractiveness to pollinators of six aromatic perennial plants from the Lamiaceae family, native and frequent in the Mediterranean region, were evaluated. Preliminary plant selection was based on designing a functional mixed margins consisting of plants attractive to pollinators and with different blooming periods, in order to extend the availability of floral resources in the field. After a year of vegetative growth, the next two years the plant species were studied in a randomized block design experiment in order to estimate their attractiveness to pollinators in Central Spain and to investigate whether floral morphology and density affect attractiveness to pollinators. The final aim of the study was to evaluate how their phenology and attractiveness to pollinators can affect the functionality of a flowering mixture of these plants. In addition, the spatial expansion, i.e. potential weediness, of the selected plant species was estimated under field conditions, as the final purpose of the studied plants is to be implemented within agro-ecosystems. The results of the experiment showed that floral morphology did not affect the attractiveness of plants to pollinators, but floral density did, as plant species with higher floral density (i.e. Nepeta tuberosa and Hyssopus officinalis) showed significantly higher attractiveness to pollinators. In addition, of six plant species, two summer species (Melissa officinalis and Thymbra capitata) did not efficiently contribute to the attractiveness of the mixture to pollinators, which reduced its attractiveness during the summer period. Finally, as none of the plants showed weedy behaviour under field conditions, the attractive plant species, i.e. N. tuberosa, H. officinalis and the early spring flowering Salvia verbenaca, showed good potential to conserve the pollinators. Similarly, in a second approximation, the attractiveness to pollinators and agronomic behaviour of twelve herbaceous plants blooming in spring were studied. This experiment was conducted over two years in a randomized block design in order to evaluate attractiveness of preselected plant species to pollinators, as well as their attractiveness efficiency (a combination of duration of flowering and insect visitation), their response to two different agronomic management practices (growing in mixed vs. mono-specific plots; tillage vs. no-tillage), and their potential weediness. The results of this experiment showed that the flowers of Borago officinalis, Echium plantagineum, Phacelia tanacetifolia and Diplotaxis tenuifolia were attractive to bees, while Calendula arvensis, Coriandrum sativum, D. tenuifolia and Lobularia maritima were attractive to hoverflies. In addition, floral mixture resulted in lower attractiveness efficiency to pollinators than mono-specific D. tenuifolia, but higher than most of the mono-specific stands. On the other hand, although some of the most attractive plant species (e.g. P. tanacetifolia and C. arvensis) showed potential weediness, their self-seeding was reduced by tillage. After comparing attractiveness efficiency of various herbaceous species to pollinators and their potential weediness, the results indicated that D. tenuifolia showed the highest attractiveness efficiency to pollinators and efficient self-reproduction, making it highly recommended to attract bees and hoverflies in agro-ecosystems of Central Spain. In addition, this plant, commonly known as wild rocket, has a supplementary economic value as a commercialized crop. The implementation of a new floral margin in agro-ecosystems means increased production costs, especially in regions with frequent and long droughts (as it is Central and South-East area of Iberian Peninsula), where the principal agricultural cost is irrigation. Therefore, before recommending D. tenuifolia for sustainable habitat management within agro-ecosystems, it is necessary to study the effect of drought stress and moderate and severe deficit irrigation on its growth, flower development and attractiveness to pollinators. The results of this experiment showed that in greenhouse conditions, potted D. tenuifolia could be without irrigation for 4 days without affecting its growth, flowering and attractiveness to pollinators. However, lack of irrigation for 8 days or longer significantly reduced the vegetative growth, number of open flowers, total floral area, flower diameter, corolla tube diameter and corolla tube length of D. tenuifolia. This study showed that regulated deficit irrigation can improve water use efficiency, and depending on the purpose of growing D. tenuifolia, as a crop or as a beneficial plant to attract pollinators, it can reduce water consumption by 40% to 70% without affecting its vegetative and floral development and without reducing its attractiveness to pollinators. Finally, the following experiment was developed in order to understand how habitat management can influence on the agricultural production. For this purpose, it was evaluated if the vicinity of mixed and mono-specific field margins, preselected to conserve pollinators within agro-ecosystems, can improve seed production in coriander (C. sativum). The selection of this plant species for the experiment was based on its necessity for insect pollination for production of seeds (even though some pollen can be transmitted from one flower to another by wind) and the fact that under semi-controlled field conditions established in the field it is possible to estimate its total seed production. Since D. tenuifolia is attractive for both bees and hoverflies in Central Spain, the main objective of this experiment was to estimate the impact of two different types of field margins, i.e. mono-specific margin with D. tenuifolia and mixed margin with six herbaceous species, on the seed production of potted coriander. For that reason, it was tested: i) if open pollination (control without proximate field margin and treatments with nearby mono-specific and mixed margin) increases the seed production of coriander when compared with no-pollination (covered inflorescences of coriander) under field conditions; ii) if frequency of pollinator visitation to the flowers of coriander was higher in the presence of field margins than in the control without field margin; and iii) if seed production was higher in the presence of field margins than in control plants of coriander without field margin. The results showed that the proximity of both types of floral margins (mixed and mono-specific) improved the seed quality of coriander plants, as seed weight and germination rate were higher than in control plants without field margin. Furthermore, the number of seeds produced was significantly higher in coriander plants grown near mixed margins than near mono-specific margin, probably due to an increase in pollinator visits. Since the experiment was conducted under semi-controlled field conditions, it can be concluded that pollinator visits was the main factor that biased the results, and that presence of both mixed or mono-specific (D. tenuifolia) margins can improve the production of coriander for more than 200% in small-scale gardens and, in addition, conserve the local pollinators.

Conductancia del mesófilo en especies forestales: implicaciones en la respuesta fotosintética a la disponibilidad de agua y luz

La fotosíntesis es el proceso biológico que permite la producción primaria y, por tanto, la vida en nuestro planeta. La tasa fotosintética viene determinada por la ‘maquinaria’ bioquímica y las resistencias difusivas al paso del CO2 desde la atmósfera hasta su fijación en el interior de los cloroplastos. Históricamente la mayor resistencia difusiva se ha atribuido al cierre estomático, sin embargo ahora sabemos, debido a las mejoras en las técnicas experimentales, que existe también una resistencia grande que se opone a la difusión del CO2 desde los espacios intercelulares a los lugares de carboxilación. Esta resistencia, llamada normalmente por su inversa: la conductancia del mesófilo (gm), puede ser igual o incluso superior a la resistencia debida por el cierre estomático. En la presente tesis doctoral he caracterizado la limitación que ejerce la resistencia del mesófilo a la fijación de CO2 en diversas especies forestales y en distintos momentos de su ciclo biológico. En la fase de regenerado, hemos estudiado tres situaciones ambientales relevantes en el mayor éxito de su supervivencia, que son: el déficit hídrico, su interacción con la irradiancia y el paso del crecimiento en la sombra a mayor irradiancia, como puede suceder tras la apertura de un hueco en el dosel forestal. En la fase de arbolado adulto se ha caracterizado el estado hídrico y el intercambio gaseoso en hojas desarrolladas a distinta irradiancia dentro del dosel vegetal durante tres años contrastados en pluviometría. Para cada tipo de estudio se han empleado las técnicas ecofisiológicas más pertinentes para evaluar el estado hídrico y el intercambio gaseoso. Por su complejidad y la falta de un método que permita su cuantificación directa, la gm ha sido evaluada por los métodos más usados, que son: la discriminación isotópica del carbono 13, el método de la J variable, el método de la J constante y el método de la curvatura. Los resultados más significativos permiten concluir que la limitación relativa a la fotosíntesis por la conductancia estomática, del mesófilo y bioquímica es dependiente de la localización de la hoja en el dosel forestal. Por primera vez se ha documentado que bajo estrés hídrico las hojas desarrolladas a la sombra estuvieron más limitadas por una reducción en la gm, mientras que las hojas desarrolladas a pleno sol estuvieron más limitadas por reducción mayor de la conductancia estomática (gsw). Encontramos buena conexión entre el aparato fotosintético foliar y el sistema hídrico debido al alto grado de correlación entre la conductancia hidráulica foliar aparente y la concentración de CO2 en los cloroplastos en distintas especies forestales. Además, hemos mostrado diferentes pautas de regulación del intercambio gaseoso según las particularidades ecológicas de las especies estudiadas. Tanto en brinzales crecidos de forma natural y en el arbolado adulto como en plántulas cultivadas en el invernadero la ontogenia afectó a las limitaciones de la fotosíntesis producidas por estrés hídrico, resultando que las limitaciones estomáticas fueron dominantes en hojas más jóvenes mientras que las no estomáticas en hojas más maduras. La puesta en luz supuso un gran descenso en la gm durante los días siguientes a la transferencia, siendo este efecto mayor según el grado de sombreo previo en el que se han desarrollado las hojas. La aclimatación de las hojas a la alta irradiancia estuvo ligada a las modificaciones anatómicas foliares y al estado de desarrollo de la hoja. El ratio entre la gm/gsw determinó la mayor eficiencia en el uso del agua y un menor estado oxidativo durante la fase de estrés hídrico y su posterior rehidratación, lo cual sugiere el uso de este ratio en los programas de mejora genética frente al estrés hídrico. Debido a que la mayoría de modelos de estimación de la producción primaria bruta (GPP) de un ecosistema no incluye la gm, los mismos están incurriendo en una sobreestimación del GPP particularmente bajo condiciones de estrés hídrico, porque más de la mitad de la reducción en fotosíntesis en hojas desarrolladas a la sombra se debe a la reducción en gm. Finalmente se presenta un análisis de la importancia en las estimas de la gm bajo estrés hídrico de la refijación del CO2 emitido en la mitocondria a consecuencia de la fotorrespiración y la respiración mitocondrial en luz. ABSTRACT Photosynthesis is the biological process that supports primary production and, therefore, life on our planet. Rates of photosynthesis are determined by biochemical “machinery” and the diffusive resistance to the transfer of CO2 from the atmosphere to the place of fixation within the chloroplasts. Historically the largest diffusive resistance was attributed to the stomata, although we now know via improvements in experimental techniques that there is also a large resistance from sub-stomatal cavities to sites of carboxylation. This resistance, commonly quantified as mesophyll conductance (gm), can be as large or even larger than that due to stomatal resistance. In the present PhD I have characterized the limitation exerted by the mesophyll resistance to CO2 fixation in different forest species at different stages of their life cycle. In seedlings, we studied three environmental conditions that affect plant fitness, namely, water deficit, the interaction of water deficit with irradiance, and the transfer of plants grown in the shade to higher irradiance as can occur when a gap opens in the forest canopy. At the stage of mature trees we characterized water status and gas exchange in leaves developed at different irradiance within the canopy over the course of three years that had contrasting rainfall. For each study we used the most relevant ecophysiological techniques to quantify water relations and gas exchange. Due to its complexity and the lack of a method that allows direct quantification, gm was estimated by the most commonly used methods which are: carbon isotope discrimination, the J-variable, constant J and the curvature method The most significant results suggest that the relative limitation of photosynthesis by stomata, mesophyll and biochemistry depending on the position of the leaf within the canopy. For the first time it was documented that under water stress shaded leaves were more limited by a reduction in gm, while the sun-adapted leaves were more limited by stomatal conductance (gsw). The connection between leaf photosynthetic apparatus and the hydraulic system was shown by the good correlations found between the apparent leaf hydraulic conductance and the CO2 concentration in the chloroplasts in shade- and sun-adapted leaves of several tree species. In addition, we have revealed different patterns of gas exchange regulation according to the functional ecology of the species studied. In field grown trees and greenhouse-grown seedlings ontogeny affected limitations of photosynthesis due to water stress with stomatal limitations dominating in young leaves and nonstomatal limitations in older leaves. The transfer to high light resulted in major decrease of gm during the days following the transfer and this effect was greater as higher was the shade which leaves were developed. Acclimation to high light was linked to the leaf anatomical changes and the state of leaf development. The ratio between the gm/gsw determined the greater efficiency in water use and reduced the oxidative stress during the water stress and subsequent rehydration, suggesting the use of this ratio in breeding programs aiming to increase avoidance of water stress. Because most models to estimate gross primary production (GPP) of an ecosystem do not include gm, they are incurring an overestimation of GPP particularly under conditions of water stress because more than half of An decrease in shade-developed leaves may be due to reduction in gm. Finally, we present an analysis of the importance of how estimates of gm under water stress are affected by the refixation of CO2 that is emitted from mitochondria via photorespiration and mitochondrial respiration in light.