Water relations of tree species growing on a rock outcrop in the "Parque Estadual de Itapuã", RS

Water relations of the tree species Myrsine umbellata Mart. ex A. DC., Dodonaea viscosa Jacq. and Erythroxylum argentinum O. E. Schulz, growing on a rock outcrop in the "Parque Estadual de Itapuã" (RS), were studied. Environmental (precipitation, temperature, soil water) and plant (water potential, vapor pressure deficit, stomatal conductance, transpiration, leaf specific hydraulic conductance, osmotic potential and cell wall elasticity) parameters were collected in five periods and pooled into two sets of data: wet and dry periods. Myrsine umbellata showed great stability of the plant parameters, including the maintenance of high pre-dawn (psiwpd) and mid-day (psiwmd) water potentials in the dry period (-0.48 and -1.12 MPa, respectively), suggesting the presence of a deep root system. Dodonaea viscosa and E. argentinum reached lower psiwpd (-1.41 and -1.97 MPa, respectively) and a greater degree of stomatal closure in the dry period, suggesting a shallower root system. Differential exposure to soil drought was also corroborated by differential drought effects on the whole-plant leaf specific hydraulic conductance (Gt). Correlation analysis pointed to weak correlations between psiwpd and g s. Erythroxylum argentinum was the only species to show osmotic adjustment in response to drought. It is suggested that M. umbellata has low tolerance to water deficits, adopting an avoidance behavior. The much lower values of psiw reached by D. viscosa and E. argentinum suggest a greater tolerance to drought by these species.

Mechanisms for the land/sea warming contrast exhibited by simulations of climate change

The land/sea warming contrast is a phenomenon of both equilibrium and transient simulations of climate change: large areas of the land surface at most latitudes undergo temperature changes whose amplitude is more than those of the surrounding oceans. Using idealised GCM experiments with perturbed SSTs, we show that the land/sea contrast in equilibrium simulations is associated with local feedbacks and the hydrological cycle over land, rather than with externally imposed radiative forcing. This mechanism also explains a large component of the land/sea contrast in transient simulations as well. We propose a conceptual model with three elements: (1) there is a spatially variable level in the lower troposphere at which temperature change is the same over land and sea; (2) the dependence of lapse rate on moisture and temperature causes different changes in lapse rate upon warming over land and sea, and hence a surface land/sea temperature contrast; (3) moisture convergence over land predominantly takes place at levels significantly colder than the surface; wherever moisture supply over land is limited, the increase of evaporation over land upon warming is limited, reducing the relative humidity in the boundary layer over land, and hence also enhancing the land/sea contrast. The non-linearity of the Clausius–Clapeyron relationship of saturation specific humidity to temperature is critical in (2) and (3). We examine the sensitivity of the land/sea contrast to model representations of different physical processes using a large ensemble of climate model integrations with perturbed parameters, and find that it is most sensitive to representation of large-scale cloud and stomatal closure. We discuss our results in the context of high-resolution and Earth-system modelling of climate change.

Regulation of plant growth in container-grown ornamentals through the use of controlled irrigation

The aim of this research was to determine whether shoot growth could be regulated and plant quality improved through two controlled irrigation techniques: Regulated Deficit Irrigation (RDI) or Partial Root Drying (PRD). An additional benefit of such techniques is that they would also improve the efficiency of irrigation application and reduce the volume of water used on commercial nurseries. Results from two ornamental woody plant species (Cotinus and Forsythia) demonstrated that plant quality could be significantly improved when RDI was applied at ≤ 60% of potential evapo-transpiration (ETp). Stomatal closure and reduced leaf and internode growth rates were associated with both the RDI and PRD techniques, but reduced leaf water potential was only recorded in the RDI system. Changes in xylem sap pH and ABA concentrations were correlated with changes in shoot physiology, and thought to be generated by those roots exposed to drying soil. By adopting such controlled irrigation systems on commercial holdings it is estimated that water consumption could be reduced by 50 to 90%.

Arabidopsis annexin1 mediates the radical-activated plasma membrane Ca2+ - and K+ -permeable conductance in root cells

Plant cell growth and stress signaling require Ca2+ influx through plasma membrane transport proteins that are regulated by reactive oxygen species. In root cell growth, adaptation to salinity stress, and stomatal closure, such proteins operate downstream of the plasma membrane NADPH oxidases that produce extracellular superoxide anion, a reactive oxygen species that is readily converted to extracellular hydrogen peroxide and hydroxyl radicals, OH_. In root cells, extracellular OH_ activates a plasma membrane Ca2+-permeable conductance that permits Ca2+ influx. In Arabidopsis thaliana, distribution of this conductance resembles that of annexin1 (ANN1). Annexins are membrane binding proteins that can form Ca2+-permeable conductances in vitro. Here, the Arabidopsis loss-of-function mutant for annexin1 (Atann1) was found to lack the root hair and epidermal OH_-activated Ca2+- and K+-permeable conductance. This manifests in both impaired root cell growth and ability to elevate root cell cytosolic free Ca2+ in response to OH_. An OH_-activated Ca2+ conductance is reconstituted by recombinant ANN1 in planar lipid bilayers. ANN1 therefore presents as a novel Ca2+-permeable transporter providing a molecular link between reactive oxygen species and cytosolic Ca2+ in plants.

Stable isotope and modelling evidence for CO2 as a driver of glacial–interglacial vegetation shifts in southern Africa

Atmospheric CO2 concentration is hypothesized to influence vegetation distribution via tree–grass competition, with higher CO2 concentrations favouring trees. The stable carbon isotope (δ13C) signature of vegetation is influenced by the relative importance of C4 plants (including most tropical grasses) and C3 plants (including nearly all trees), and the degree of stomatal closure – a response to aridity – in C3 plants. Compound-specific δ13C analyses of leaf-wax biomarkers in sediment cores of an offshore South Atlantic transect are used here as a record of vegetation changes in subequatorial Africa. These data suggest a large increase in C3 relative to C4 plant dominance after the Last Glacial Maximum. Using a process-based biogeography model that explicitly simulates 13C discrimination, it is shown that precipitation and temperature changes cannot explain the observed shift in δ13C values. The physiological effect of increasing CO2 concentration is decisive, altering the C3/C4 balance and bringing the simulated and observed δ13C values into line. It is concluded that CO2 concentration itself was a key agent of vegetation change in tropical southern Africa during the last glacial–interglacial transition. Two additional inferences follow. First, long-term variations in terrestrial δ13Cvalues are not simply a proxy for regional rainfall, as has sometimes been assumed. Although precipitation and temperature changes have had major effects on vegetation in many regions of the world during the period between the Last Glacial Maximum and recent times, CO2 effects must also be taken into account, especially when reconstructing changes in climate between glacial and interglacial states. Second, rising CO2 concentration today is likely to be influencing tree–grass competition in a similar way, and thus contributing to the "woody thickening" observed in savannas worldwide. This second inference points to the importance of experiments to determine how vegetation composition in savannas is likely to be influenced by the continuing rise of CO2 concentration.

Transgenerational, dynamic methylation of stomata genes in response to low relative humidity

Transgenerational inheritance of abiotic stress-induced epigenetic modifications in plants has potential adaptive significance and might condition the offspring to improve the response to the same stress, but this is at least partly dependent on the potency, penetrance and persistence of the transmitted epigenetic marks. We examined transgenerational inheritance of low Relative Humidity-induced DNA methylation for two gene loci in the stomatal developmental pathway in Arabidopsis thaliana and the abundance of associated short-interfering RNAs (siRNAs). Heritability of low humidity-induced methylation was more predictable and penetrative at one locus (SPEECHLESS, entropy ≤ 0.02; χ2 < 0.001) than the other (FAMA, entropy ≤ 0.17; χ2 ns). Methylation at SPEECHLESS correlated positively with the continued presence of local siRNAs (r2 = 0.87; p = 0.013) which, however, could be disrupted globally in the progeny under repeated stress. Transgenerational methylation and a parental low humidity-induced stomatal phenotype were heritable, but this was reversed in the progeny under repeated treatment in a previously unsuspected manner.

Desiccation tolerance in Pleurostima purpurea (Velloziaceae)

Previous works suggested that Pleurostima purpurea (Velloziaceae-Barbacenioideae) shows a remarkable capacity to endure desiccation of its vegetative tissues. P. purpurea occurs in monocotyledons mats on soil islands in the Pao de Acucar (Sugar Loaf) one of the most recognizable rock outcrops of the world, in Rio de Janeiro, southeastern Brazil. Mats of P. purpurea occur in cliffs by the sea some meters above the tidal zone. Although living in rock outcrops almost devoid of any soil cover, P. purpurea seems to occur preferably on less exposed rock faces and slightly shady sites. Usually, less extreme adaptations to drought would be expected in plants with the habitat preference of P. purpurea. Relying on this observation, we argue if a combination of different strategies of dealing with low water availability can be found in P. purpurea as on other desiccation tolerant angiosperms. This study aims to examine the occurrence of desiccation tolerant behavior in P. purpurea together with the expression of drought avoidance mechanisms during dehydration progression. For this, it was analyzed the gas exchanges, leaf pigments and relative leaf water content during desiccation and rehydration of cultivated mature individuals. P. purpurea behaved like typical drought avoiders under moderated drought condition with stomatal closure occurring around a relative leaf water content up to 90%. During this process, it was observed a delay in the leaf relative water content (RWC(leaf)) decrease comparing to the plant-soil relative water content (RWC(plant-soil)). As soil dehydration worsened, gas exchanges restrictions progressed until a lack of activity which characterizes anabiosis. The loss of chlorophyll occurs before the end of total dehydration, characterizing the presence of poikilochlorophylly. The chlorophyll degradation follows the RWC(leaf) decrease, which achieved the minimum average value of 17% without incurring in leaf abscission. The chlorophyll re-synthesis seems to start well after the full rehydration of the leaf. During all of this process, carotenoid content remained stable. These results are coherent with a combination of drought avoidance and desiccation tolerance in P. purpurea which seems to be coherent with the amplitude of water availability in the rock outcrop habitat where it occurs, suggesting that the periods of water availability are sufficiently long for the success of the costly desiccation tolerant behavior but too short to make a typical drought avoider species win the competition for exploring the rock outcrop substrate where P. purpurea occurs.

An Arabidopsis Mitochondrial Uncoupling Protein Confers Tolerance to Drought and Salt Stress in Transgenic Tobacco Plants

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

Photosynthetic response of soybean to twospotted spider mite (Acari: Tetranychydae) injury

Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES)

Variação sazonal da fotossíntese, condutância estomática e potencial da água na folha de laranjeira 'Valência'

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Effects of potassium and sodium supply on drought-adaptive mechanisms in Eucalyptus grandis plantations

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Overexpression of the activated form of the AtAREB1 gene (AtAREB1 Delta QT) improves soybean responses to water deficit

Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES)

Cultivo de Eucalyptus urograndis em atmosfera enriquecida com CO2: mudanças no proteoma cloroplastidial

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

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.

Respuestas anatómico fisiológicas frente a estrés hídrico en plantaciones de especies de crecimiento rápido para la producción de biomasa

El objetivo general de la presente Tesis es identificar algunas de las características anatómico-fisiológicas que confieren la capacidad de alcanzar una mejor productividad bajo clima mediterráneo a plantas de diversos genotipos de los géneros Populus y Eucalyptus, caracterizados por su carácter pionero, elevado crecimiento y vulnerabilidad a la cavitación. En los dos primeros capítulos se hace un seguimiento de la conductancia estomática a una selección de clones de eucalipto cultivados en invernadero, sometidos a diferentes dotaciones hídricas. Se realizaron además mediciones periódicas del pH de la savia del tallo y de la pérdida de conductividad hidráulica para investigar su implicación en la regulación química e hidraúlica del cierre estomático. Las variaciones en el pH de la savia obtenidas parecen responder a cambios en el déficit de presión de vapor de agua atmosférico y no a diferencias en la disponibilidad de agua en el suelo. La conductancia estomática presentó una correlación positiva significativa con el pH de la savia, pero no con la conductividad hidráulica. La variabilidad de la conductividad hidráulica máxima se discute a la luz de recientes investigaciones sobre los materiales constituyentes de las membranas de las punteaduras. Los clones que mostraron mayores conductancias hidráulicas y estomáticas presentaron valores más altos de producción y supervivencia, poniendo de manifiesto la utilidad del estudio de estas variables. Por el contrario, los valores más bajos de conductancia estomática e hidraúlica se encontraron en clones que han resultado un fracaso en plantaciones comerciales, en particular, fue destacable el mal resultado de un clon procedente de autocruzamiento respecto de otros híbridos. En el tercer capítulo de la tesis se estudian características anatómicas y funcionales del xilema relacionadas con la eficiencia en el transporte de agua a las hojas, y que pueden afectar directa o indirectamente a la transpiración y al crecimiento. Los estudios anatómicos fueron realizados sobre brotes anuales de chopo en una plantación situada en Granada, en condiciones de riego limitante. La combinación de rasgos anatómicos más favorable de cara a la producción de biomasa fue una densidad alta de vasos de diámetro intermedio. Los clones más productivos figuraron entre los más resistentes a la cavitación. Para estudiar el crecimiento de masas arboladas se utilizan frecuentemente parámetros fisiológicos como el índice de area foliar (LAI). La estimación del LAI a partir de fotografías hemisféricas aplicada a tallares de chopo plantados a alta densidad y a turno corto para biomasa se lleva a cabo mediante una metodología reciente empleada y discutida en el cuarto capítulo de la Tesis. Los resultados muestran que las diferencias de producción existentes entre genotipos, localidades de medición con diferentes dosis de riego, y años, pueden predecirse a partir de la determinación del índice de área foliar tanto por métodos directos como indirectos de estimación. Tanto los estudios realizados en eucalipto como en chopo han mostrado que los genotipos con menores producciones de biomasa en campo alcanzaron los menores valores de conductancia estomática en las condiciones más favorables así como el menor número de vasos en el xilema. La estrecha relación entre crecimiento y LAI confirma una vez más la importancia del desarrollo de la copa para sostener un buen crecimiento. El mayor desarrollo de la copa y rendimiento en biomasa se midieron en uno de los clones con un número de vasos más elevado, y menor vulnerabilidad a la cavitación en condiciones de estrés. Estos resultados ponen de manifiesto la importancia de las características anatómicas y funcionales del xilema como condicionantes del patrón de crecimiento de las plantas y el comportamiento de los estomas. ABSTRACT A number of anatomical xylem traits and physiological variables were analyzed in genotypes of both the Populus and Eucalyptus genera with the main aim of identifying traits in the genotypes which confer the ability to produce an acceptable biomass yield under Mediterranean climatic conditions. In the first two chapters of this PhD, the results of two experiments carried out on several clones of the species Eucalyptus globulus Labill. are presented. Chapters three and four include the results of another two trials on four poplar hybrid genotypes. One of the initial plant responses to water stress is stomatal closure, which can be triggered by hydraulic and/or chemical signals. The two first chapters of this PhD deal with trials in which stomatal conductance and percentage loss of hydraulic conductivity were monitored on a set of eucalyptus clones supplied by ENCE (former National Cellulose Company) and currently used in the company’s own commercial plantings. The experimental trials were carried out in greenhouses and the plants were submitted to two different watering regimes. The pH of the stem sap was periodically measured as the greenhouse temperature and humidity changed. The aim of these measurements was to investigate the role of both sap pH and percentage loss of hydraulic conductivity on stomatal regulation. The results obtained suggest that changes in sap pH are a response to vapor pressure deficit changes rather than to differences in soil water availability. We found significant correlation between stomatal conductance and sap pH, although no significant relationship was found between stomatal conductance and hydraulic conductivity. Variability in maximum hydraulic conductivity is discussed based on recent pit membrane constituent research. The study of hydraulic conductivity proved helpful in order to detect the clones with both higher growth and greater chance of survival, since clones displaying the lowest hydraulic conductivities were those that failed in commercial plantings. Anatomical xylem traits define the water transport efficiency to leaves and can therefore limit transpiration and growth. The third chapter of this PhD addresses anatomical xylem traits in poplar. One year old stem samples were taken from a water-stressed trial in Granada. The anatomical xylem study proved useful for detecting the lowest yielding genotypes. Clones with intermediate vessel size and high vessel densities were found to be those with the highest biomass yield. Differences in cavitation resistance depending on the clone tested and the water treatment applied were also found. The clones with the highest biomass yield were found to be among the most cavitation resistant clones in each watering regime. Xylem and physiological traits along with stomatal behavior are useful tools to determine plant growth. In order to study plantings or forests, it is more common to employ other physiological variables such as leaf area index (LAI). LAI estimation from hemispherical photographs applied to short rotation woody crops is a recently developed method that still requires fine tuning through further investigation. In the fourth chapter, data from LAI monitoring over two consecutive years were analyzed in two different locations where different irrigation treatments were applied. The results showed that differences in yield between genotypes, different irrigation regimes and years could be predicted by using the LAI estimates, either through direct or indirect estimation methods. Our studies of poplar and eucalyptus have shown that the field-grown genotypes with the lowest biomass yield displayed the lowest values of stomatal conductance under the most favorable environmental conditions and also had a low number of xylem conduits. The close relationship between LAI and growth highlights the importance of crown development in biomass growth. The highest LAI and biomass yield were recorded in one of the clones with higher vessel density and the lowest vulnerability to cavitation under stress conditions. These results underline the importance of research into anatomical and functional traits as factors influencing plant growth patterns and stomatal behavior.