Species-specific stomatal response of trees to drought - a link to vegetation dynamics?

Question: Is stomatal regulation specific for climate and tree species, and does it reveal species-specific responses to drought? Is there a link to vegetation dynamics? Location: Dry inner alpine valley, Switzerland Methods: Stomatal aperture (θE) of Pinus sylvestris, Quercus pubescens, Juniperus communis and Picea abies were continuously estimated by the ratio of measured branch sap flow rates to potential transpiration rates (adapted Penman-Monteith single leaf approach) at 10-min intervals over four seasons. Results: θE proved to be specific for climate and species and revealed distinctly different drought responses: Pinus stomata close disproportionately more than neighbouring species under dry conditions, but has a higher θE than the other species when weather was relatively wet and cool. Quercus keeps stomata more open under drought stress but has a lower θE under humid conditions. Juniperus was most drought-tolerant, whereas Picea stomata close almost completely during summer. Conclusions: The distinct microclimatic preferences of the four tree species in terms of θE strongly suggest that climate (change) is altering tree physiological performances and thus species-specific competitiveness. Picea and Pinus currently live at the physiological limit of their ability to withstand increasing temperature and drought intensities at the sites investigated, whereas Quercus and Juniperus perform distinctly better. This corresponds, at least partially, with regional vegetation dynamics: Pinus has strongly declined, whereas Quercus has significantly increased in abundance in the past 30 years. We conclude that θE provides an indication of a species' ability to cope with current and predicted climate.

Selection and hydroponic growth of bread wheat cultivars for bioregenerative life support systems

As part of the ESA-funded MELiSSA program, the suitability, the growth and the development of four bread wheat cultivars were investigated in hydroponic culture with the aim to incorporate such a cultivation system in an Environmental Control and Life Support System (ECLSS). Wheat plants can fulfill three major functions in space: (a) fixation of CO2 and production of O2, (b) production of grains for human nutrition and (c) production of cleaned water after condensation of the water vapor released from the plants by transpiration. Four spring wheat cultivars (Aletsch, Fiorina, Greina and CH Rubli) were grown hydroponically and compared with respect to growth and grain maturation properties. The height of the plants, the culture duration from germination to harvest, the quantity of water used, the number of fertile and non-fertile tillers as well as the quantity and quality of the grains harvested were considered. Mature grains could be harvested after around 160 days depending on the varieties. It became evident that the nutrient supply is crucial in this context and strongly affects leaf senescence and grain maturation. After a first experiment, the culture conditions were improved for the second experiment (stepwise decrease of EC after flowering, pH adjusted twice a week, less plants per m2) leading to a more favorable harvest (higher grain yield and harvest index). Considerably less green tillers without mature grains were present at harvest time in experiment 2 than in experiment 1. The harvest index for dry matter (including roots) ranged from 0.13 to 0.35 in experiment 1 and from 0.23 to 0.41 in experiment 2 with modified culture conditions. The thousand-grain weight for the four varieties ranged from 30.4 to 36.7 g in experiment 1 and from 33.2 to 39.1 g in experiment 2, while market samples were in the range of 39.4–46.9 g. Calcium levels in grains of the hydroponically grown wheat were similar to those from field-grown wheat, while potassium, magnesium, phosphorus, iron, zinc, copper, manganese and nickel levels tended to be higher in the grains of experimental plants. It remains a challenge for future experiments to further adapt the nutrient supply in order to improve senescence of vegetative plant parts, harvest index and the composition of bread wheat grains.

Tree species and diversity effects on soil water seepage in a tropical plantation

Plant diversity has been shown to influence the water cycle of forest ecosystems by differences in water consumption and the associated effects on groundwater recharge. However, the effects of biodiversity on soil water fluxes remain poorly understood for native tree species plantations in the tropics. Therefore, we estimated soil water fluxes and assessed the effects of tree species and diversity on these fluxes in an experimental native tree species plantation in Sardinilla (Panama). The study was conducted during the wet season 2008 on plots of monocultures and mixtures of three or six tree species. Rainfall and soil water content were measured and evapotranspiration was estimated with the Penman-Monteith equation. Soil water fluxes were estimated using a simple soil water budget model considering water input, output, and soil water and groundwater storage changes and in addition, were simulated using the physically based one-dimensional water flow model Hydrus-1D. In general, the Hydrus simulation did not reflect the observed pressure heads, in that modeled pressure heads were higher compared to measured ones. On the other hand, the results of the water balance equation (WBE) reproduced observed water use patterns well. In monocultures, the downward fluxes through the 200 cm-depth plane were highest below Hura crepitans (6.13 mm day−1) and lowest below Luehea seemannii (5.18 mm day−1). The average seepage rate in monocultures (±SE) was 5.66 ± 0.18 mm day−1, and therefore, significantly higher than below six-species mixtures (5.49 ± 0.04 mm day−1) according to overyielding analyses. The three-species mixtures had an average seepage rate of 5.63 ± 0.12 mm day−1 and their values did not differ significantly from the average values of the corresponding species in monocultures. Seepage rates were driven by the transpiration of the varying biomass among the plots (r = 0.61, p = 0.017). Thus, a mixture of trees with different growth rates resulted in moderate seepage rates compared to monocultures of either fast growing or slow growing tree species. Our results demonstrate that tree-species specific biomass production and tree diversity are important controls of seepage rates in the Sardinilla plantation during the wet season.

Stomatal opening at elevated temperature: an underestimated regulatory mechanism?

Climate models predict more frequent and more severe extreme events (e.g. heat waves, extended drought periods) in Europe during the next decades. The response of plants to elevated temperature is a key issue in this context. Stomatal regulation is not only relevant for the diffusion of CO2 from the ambient air into the leaves, but it plays also an important role for the control of transpiration and leaf cooling. The regulation of stomatal aperture by the water status (hydroactive and hydropassive feed-back) and by internal CO2 availability (CO2 feed-back) are well documented in the literature, while the response of the stomates to elevated temperature was far less considered in the past. Photosynthesis is negatively affected by elevated temperature, but the water loss via transpiration may still be high. In the experiments reported here, bean leaf segments were incubated in darkness floating on water in the range from 20 to 50°C and then analyzed immediately by taking a photograph with a digital microscope. Stomatal aperture was measured on these pictures in order to quantify stomatal opening. After the incubation for 30 min, the opening was 0.66, 2.76 and 4.28 μm at 23, 30 and 35°C respectively. This opening at elevated temperature was fully reversible. Abscisic acid (0.1 μM) in the incubation medium shifted the temperature for stomatal opening to higher values. It can be concluded that elevated temperature stimulates stomatal opening regardless of the CO2 assimilation status and that there is a trade-off between leaf cooling on one hand and limiting water loss during drought periods on the other hand.

Extreme climatic events: impacts of drought and high temperature on physiological processes in agronomically important plants

Climate models predict more frequent and more severe extreme events (e.g., heat waves, extended drought periods, flooding) in many regions for the next decades. The impact of adverse environmental conditions on crop plants is ecologically and economically relevant. This review is focused on drought and heat effects on physiological status and productivity of agronomically important plants. Stomatal opening represents an important regulatory mechanism during drought and heat stress since it influences simultaneously water loss via transpiration and CO2 diffusion into the leaf apoplast which further is utilized in photosynthesis. Along with the reversible short-term control of stomatal opening, stomata and leaf epidermis may produce waxy deposits and irreversibly down-regulate the stomatal conductance and non-stomatal transpiration. As a consequence photosynthesis will be negatively affected. Rubisco activase—a key enzyme in keeping the Calvin cycle functional—is heat-sensitive and may become a limiting factor at elevated temperature. The accumulated reactive oxygen species (ROS) during stress represent an additional challenge under unfavorable conditions. Drought and heat cause accumulation of free amino acids which are partially converted into compatible solutes such as proline. This is accompanied by lower rates of both nitrate reduction and de novo amino acid biosynthesis. Protective proteins (e.g., dehydrins, chaperones, antioxidant enzymes or the key enzyme for proline biosynthesis) play an important role in leaves and may be present at higher levels under water deprivation or high temperatures. On the whole plant level, effects on long-distance translocation of solutes via xylem and phloem and on leaf senescence (e.g., anticipated, accelerated or delayed senescence) are important. The factors mentioned above are relevant for the overall performance of crops under drought and heat and must be considered for genotype selection and breeding programs.

Pasture degradation modifies the water and carbon cycles of the Tibetan highlands

The Tibetan Plateau has a significant role with regard to atmospheric circulation and the monsoon in particular. Changes between a closed plant cover and open bare soil are one of the striking effects of land use degradation observed with unsustainable range management or climate change, but experiments investigating changes of surface properties and processes together with atmospheric feedbacks are rare and have not been undertaken in the world's two largest alpine ecosystems, the alpine steppe and the Kobresia pygmaea pastures of the Tibetan Plateau. We connected measurements of micro-lysimeter, chamber, 13C labelling, and eddy covariance and combined the observations with land surface and atmospheric models, adapted to the highland conditions. This allowed us to analyse how three degradation stages affect the water and carbon cycle of pastures on the landscape scale within the core region of the Kobresia pygmaea ecosystem. The study revealed that increasing degradation of the Kobresia turf affects carbon allocation and strongly reduces the carbon uptake, compromising the function of Kobresia pastures as a carbon sink. Pasture degradation leads to a shift from transpiration to evaporation while a change in the sum of evapotranspiration over a longer period cannot be confirmed. The results show an earlier onset of convection and cloud generation, likely triggered by a shift in evapotranspiration timing when dominated by evaporation. Consequently, precipitation starts earlier and clouds decrease the incoming solar radiation. In summary, the changes in surface properties by pasture degradation found on the highland have a significant influence on larger scales.

Heavy Metals in Crop Plants: Transport and Redistribution Processes on the Whole Plant Level

Copper, zinc, manganese, iron, nickel and molybdenum are essential micronutrients for plants. However, when present in excess they may damage the plant or decrease the quality of harvested plant products. Some other heavy metals such as cadmium, lead or mercury are not needed by plants and represent pollutants. The uptake into the roots, the loading into the xylem, the acropetal transport to the shoot with the transpiration stream and the further redistribution in the phloem are crucial for the distribution in aerial plant parts. This review is focused on long-distance transport of heavy metals via xylem and phloem and on interactions between the two transport systems. Phloem transport is the basis for the redistribution within the shoot and for the accumulation in fruits and seeds. Solutes may be transferred from the xylem to the phloem (e.g., in the small bundles in stems of cereals, in minor leaf veins). Nickel is highly phloem-mobile and directed to expanding plant parts. Zinc and to a lesser degree also cadmium are also mobile in the phloem and accumulate in meristems (root tips, shoot apex, axillary buds). Iron and manganese are characterized by poor phloem mobility and are retained in older leaves.

Simulated heat waves affected alpine grassland only in combination with drought

The Alpine region is warming fast, and concurrently, the frequency and intensity of climate extremes are increasing. It is currently unclear whether alpine ecosystems are sensitive or resistant to such extremes. We subjected Swiss alpine grassland communities to heat waves with varying intensity by transplanting monoliths to four different elevations (2440–660 m above sea level) for 17 d. Half of these were regularly irrigated while the other half were deprived of irrigation to additionally induce a drought at each site. Heat waves had no significant impacts on fluorescence (Fv/Fm, a stress indicator), senescence and aboveground productivity if irrigation was provided. However, when heat waves coincided with drought, the plants showed clear signs of stress, resulting in vegetation browning and reduced phytomass production. This likely resulted from direct drought effects, but also, as measurements of stomatal conductance and canopy temperatures suggest, from increased high-temperature stress as water scarcity decreased heat mitigation through transpiration. The immediate responses to heat waves (with or without droughts) recorded in these alpine grasslands were similar to those observed in the more extensively studied grasslands from temperate climates. Responses following climate extremes may differ in alpine environments, however, because the short growing season likely constrains recovery.

Long-distance transport of alkali metals in maturing wheat

The alkali metals cesium, rubidium, lithium and sodium were introduced together with strontium via flaps into leaf laminas or into the stem of maturing, intact winter wheat (Triticum aestivum L. cv. Arina) grown in a field. Long-distance transport of these elements and the influence of the application date and of different application positions were investigated. The phloem-immobile Sr served as a marker for the distribution of the xylem sap in the plants. Dry matter accumulation in the grains and the transpiration per shoot were not markedly affected by the treatments as compared to control plants. The phloem mobility was rather high for Cs and Rb. Li was almost immobile in the phloem (similarly to Sr). An application into the cut stem xylem below the second leaf node contributed more to the contents in the grains than an application into the flag leaf. An earlier feeding date led to a higher accumulation in the grains. The marked losses of the elements applied during maturation (most pronounced for Li) can be explained by leakage in the rain.

Relaciones entre cavitación y cierre estomático en vides sometidas a déficit hídrico

La cavitación es una disfunción fisiológica que ocurre en el xilema de las plantas cuando estas están bajo déficit hídrico, y que entraña una pérdida de su conductancia hidráulica (kL), cuando algunos vasos se llenan de aire. Esto incide negativamente sobre la oferta de agua y afecta el potencial hídrico foliar (ΨL) y la hidratación de la canopia. El cierre estomático es una respuesta efectiva ante la disminución del contenido hídrico. Dependiendo de la especie vegetal, los estomas suelen cerrase para evitar la cavitación catastrófica. Una planta poco vulnerable a cavitar puede mantenerlos abiertos por más tiempo y por lo tanto seguir fotosintetizando. Por el contrario, plantas vulnerables a la cavitación deben cerrar sus estomas con anterioridad para poder evitar cavitaciones graves. En este trabajo se estudió, el ajuste estomático como mecanismo para evitar la cavitación en cuatro variedades contrastantes de vid (Grenache, Syrah, Malbec y Chardonnay). Se hipotetizó que las vides sometidas a déficit hídrico disminuyen gs para evitar sufrir cavitaciones catastróficas y que algunas variedades de vid, cuando crecen bajo restricción hídrica, se aclimatan mejor desarrollando un ajuste estomático más preciso, ó un sistema conductor menos vulnerable a cavitar, o ambas. Se diseñó un experimento aleatorizado en invernáculo donde se probaron dos situaciones hídricas edáficas (100% y 50% de FTSW). Luego de dos meses, se midieron, a lo largo del día, las variables gs, transpiración y potencial hídrico. Luego se construyeron las curvas de cavitación y se calculó la conduactancia hidráulica de la planta (kL) y el embolismo a lo largo del día. Finalmente se obtuvo la relación entre gs, kL y el embolismo. Mediante un modelo mecanístico, que se construyó teniendo en cuenta los flujos de agua y vapor, las kL y gs, y la vulnerabilidad del xilema a cavitar; se probó que gs no es la única variable responsable de frenar la embolia. Se determinó que gs y kL están íntimamente asociadas y que este acople entre ambas conductancias es lo que frena la embolia. Se concluyó que, en la vid y bajo niveles de estrés hídrico moderado, no es necesario un cierre estomático para controlar la cavitación, sino una disminución de la diferencia entre gs y kL (Δgs). Por esto, el mecanismo de control de la cavitación en la vid no conlleva un costo en términos de intercambio gaseoso. También se descubrió que bajo niveles moderados de déficit hídrico la vulnerabilidad xilemática no aumenta con respecto a las plantas que no sufren déficit, sin importar de qué variedad se trate.

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.

Cantidad de agua que transpira la vid (Vitis vinifera L.) : variedad cultígena "semillón" : en Mendoza

Se realizaron numerosas observaciones de transpiración vegetal con vid (Vitis vinifera L.), variedad cultígena “Semillón", desde la brotación en primavera hasta la caída de las hojas en otoño. El fin principal del estudio fué poder determinar la cantidad de agua que una hectárea de cultivo de vid transpira en Mendoza (Argentina), durante su ciclo biológico completo, y que fue de 4.243,5 m³. Las experiencias se realizaron en los campos experimentales y _ laboratorios de la Facultad de Ciencias Agrarias de la Universidad Nacional de Cuyo. El viñedo estudiado era de tipo espaldera baja, con plantas de vid en hileras distanciadas 2 m y habiendo 1 m de distancia entre planta y planta, lo que suma 5.000 cepas por hectárea. Se consideró un período activo de transpiración de 207 días. La transpiración media diaria (24 horas) por planta fue de 4,1 litros de agua.

Model results in NetCDF format of North African vegetation-precipitation in early and mid-Holocene climate simulated by CCSM3-DGVM//

The present study analyses the sign, strength, and working mechanism of the vegetation-precipitation feedback over North Africa in middle (6 ka BP) and early Holocene (9 ka BP) simulations using the comprehensive coupled climate-vegetation model CCSM3-DGVM (Community Climate System Model version 3 and a dynamic global vegetation model). The coupled model simulates enhanced summer rainfall and a northward migration of the West African monsoon trough along with an expansion of the vegetation cover for the early and middle Holocene compared to the pre-industrial period. It is shown that dynamic vegetation enhances the orbitally triggered summer precipitation anomaly by approximately 20% in the Sahara-Sahel region (10-25° N, 20° W-30° E) in both the early and mid-Holocene experiments compared to their fixed-vegetation counterparts. The primary vegetation-rainfall feedback identified here operates through surface latent heat flux anomalies by canopy evaporation and transpiration and their effect on the mid-tropospheric African easterly jet, whereas the effects of vegetation changes on surface albedo and local water recycling play a negligible role. Even though CCSM3-DGVM simulates a positive vegetation-precipitation feedback in the North African region, this feedback is not strong enough to produce multiple equilibrium climate-ecosystem states on a regional scale.

Fractal Behaviour and Irregularity of Wetting Fronts in Heterogeneous Porous Media

The wetting front is the zone where water invades and advances into an initially dry porous material and it plays a crucial role in solute transport through the unsaturated zone. Water is an essential part of the physiological process of all plants. Through water, necessary minerals are moved from the roots to the parts of the plants that require them. Water moves chemicals from one part of the plant to another. It is also required for photosynthesis, for metabolism and for transpiration. The leaching of chemicals by wetting fronts is influenced by two major factors, namely: the irregularity of the fronts and heterogeneity in the distribution of chemicals, both of which have been described by using fractal techniques. Soil structure can significantly modify infiltration rates and flow pathways in soils. Relations between features of soil structure and features of infiltration could be elucidated from the velocities and the structure of wetting fronts. When rainwater falls onto soil, it doesn?t just pool on surfaces. Water ?or another fluid- acts differently on porous surfaces. If the surface is permeable (porous) it seeps down through layers of soil, filling that layer to capacity. Once that layer is filled, it moves down into the next layer. In sandy soil, water moves quickly, while it moves much slower through clay soil. The movement of water through soil layers is called the the wetting front. Our research concerns the motion of a liquid into an initially dry porous medium. Our work presents a theoretical framework for studying the physical interplay between a stationary wetting front of fractal dimension D with different porous materials. The aim was to model the mass geometry interplay by using the fractal dimension D of a stationary wetting front. The plane corresponding to the image is divided in several squares (the minimum correspond to the pixel size) of size length ". We acknowledge the help of Prof. M. García Velarde and the facilities offered by the Pluri-Disciplinary Institute of the Complutense University of Madrid. We also acknowledge the help of European Community under project Multi-scale complex fluid flows and interfacial phenomena (PITN-GA-2008-214919). Thanks are also due to ERCOFTAC (PELNoT, SIG 14)

Importance of canopy porosity into vineyard and the relationship with the grape maturity. Digital estimation method

In warm and dry climates, the use of porous systems should be required in order to allow a better leaf distribution inside the plant, causing more space in the clusters area and enhancing determined physiological processes so in the leaf (photosynthesis, v entilation, transpiration) as in berry (growth and maturation). Plant geometry indexes, yield and must composition have been studied in three different systems: sprawl with 12 shoots/m (S1); sprawl system with 18 shoots/m (S2) and vertical positioned syste m or VSP with 12 shoots/m (VSP1). Total leaf area increases as the crop load does, whoever surface area depends on to two factors: crop load and the training system (VSP vs. sprawl), which can provide differences in leaf exposure efficiencies. The main objective of this study was to validate digital photography measurements used to compare porosity differences among treatments and, as they affect plant microclimate and, therefore, yield and berry quality. Also, all previous studied indexes (LAI, SA, SFEr) tended to overestimate the relationship between exposed leaf surface and porosity of each treatment, but the use of digital method proved to be an effective tool in order to assess canopy porosity. Results showed that not positioned and free systems (sprawl) scored between 25- 50% more porosity in the clusters area than the fixed vertical system (VSP), which resulted in a better plant microclimate for test conditions, mainly by improving the exposure of internal clusters and internal canopy ventilation. On the other hand, higher crop load treatment (S2) showed a real increase in yield (16%) without any relevant change into must composition, even improving total anthocyanin content into berry during ripening