Water-use efficiency and transpiration across European forests during the Anthropocene

The Earth’s carbon and hydrologic cycles are intimately coupled by gas exchange through plant stomata1, 2, 3. However, uncertainties in the magnitude4, 5, 6 and consequences7, 8 of the physiological responses9, 10 of plants to elevated CO2 in natural environments hinders modelling of terrestrial water cycling and carbon storage11. Here we use annually resolved long-term δ13C tree-ring measurements across a European forest network to reconstruct the physiologically driven response of intercellular CO2 (Ci) caused by atmospheric CO2 (Ca) trends. When removing meteorological signals from the δ13C measurements, we find that trees across Europe regulated gas exchange so that for one ppmv atmospheric CO2 increase, Ci increased by ~0.76 ppmv, most consistent with moderate control towards a constant Ci/Ca ratio. This response corresponds to twentieth-century intrinsic water-use efficiency (iWUE) increases of 14 ± 10 and 22 ± 6% at broadleaf and coniferous sites, respectively. An ensemble of process-based global vegetation models shows similar CO2 effects on iWUE trends. Yet, when operating these models with climate drivers reintroduced, despite decreased stomatal opening, 5% increases in European forest transpiration are calculated over the twentieth century. This counterintuitive result arises from lengthened growing seasons, enhanced evaporative demand in a warming climate, and increased leaf area, which together oppose effects of CO2-induced stomatal closure. Our study questions changes to the hydrological cycle, such as reductions in transpiration and air humidity, hypothesized to result from plant responses to anthropogenic emissions.

Intra-annual radial growth and water relations of trees - implications towards a growth mechanism

There is a missing link between tree physiological and wood-anatomical knowledge which makes it impossible mechanistically to explain and predict the radial growth of individual trees from climate data. Empirical data of microclimatic factors, intra-annual growth rates, and tree-specific ratios between actual and potential transpiration (T PET−1) of trees of three species (Quercus pubescens, Pinus sylvestris, and Picea abies) at two dry sites in the central Wallis, Switzerland, were recorded from 2002 to 2004 at a 10 min resolution. This included the exceptionally hot and dry summer of 2003. These data were analysed in terms of direct (current conditions) and indirect impacts (predispositions of the past year) on growth. Rain was found to be the only factor which, to a large extent, consistently explained the radial increment for all three tree species at both sites and in the short term as well. Other factors had some explanatory power on the seasonal time-scale only. Quercus pubescens built up much of its tree ring before bud break. Pinus sylvestris and Picea abies started radial growth 1–2 weeks after Quercus pubescens and this was despite the fact that they had a high T PET−1 before budburst and radial growth started. A high T PET−1 was assumed to be related to open stomata, a very high net CO2 assimilation rate, and thus a potential carbon (C)-income for the tree. The main period of radial growth covered about 30–70% of the productive days of a year. In terms of C-allocation, these results mean that Quercus pubescens depended entirely on internal C-stores in the early phase of radial growth and that for all three species there was a long time period of C-assimilation which was not used for radial growth in above-ground wood. The results further suggest a strong dependence of radial growth on the current tree water relations and only secondarily on the C-balance. A concept is discussed which links radial growth over a feedback loop to actual tree water-relations and long-term affected C-storage to microclimate.

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.

Effect of locally increased zinc contents on zinc transport from the flag leaf lamina to the maturing grains of wheat

From previous experiments, it was evident that the accumulation of zinc in maturing wheat grains is highly regulated, but the regulatory mechanisms involved are not yet identified. In this study, we determined the transfer of radiolabelled zinc (fed directly into a leaf flap) from the flag leaf lamina to the grains. We also determined how this zinc transfer was affected by feeding additional unlabeled zinc (1 μmol per plant) either into the flag leaf sheath or the peduncle. Most of the 65Zn was retained in the feeding flap. A high percentage of the zinc exported from the flap accumulated in the grains with little accumulation of radiolabel in the other parts of the shoot. Unlabeled zinc remained mainly in the feeding flap and in the parts reached by the transpiration stream from the feeding position. The transfer of radiolabelled zinc was essentially not influenced by unlabeled zinc fed into another plant part. Our results suggest that the loading of zinc into the phloem and the mass flow in the sieve tubes might regulate zinc redistribution within the wheat shoot.