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.

Growth at moderately elevated temperature alters the physiological response of the photosynthetic apparatus to heat stress in pea (Pisum sativum L.) leaves

The impact of heat stress on the functioning of the photosynthetic apparatus was examined in pea (Pisum sativum L.) plants grown at control (25 °C; 25 °C-plants) or moderately elevated temperature (35 °C; 35 °C-plants). In both types of plants net photosynthesis (Pn) decreased with increasing leaf temperature (LT) and was more than 80% reduced at 45 °C as compared to 25 °C. In the 25 °C-plants, LTs higher than 40 °C could result in a complete suppression of Pn. Short-term acclimation to heat stress did not alter the temperature response of Pn. Chlorophyll a fluorescence measurements revealed that photosynthetic electron transport (PET) started to decrease when LT increased above 35 °C and that growth at 35 °C improved the thermal stability of the thylakoid membranes. In the 25 °C-plants, but not in the 35 °C-plants, the maximum quantum yield of the photosystem II primary photochemistry, as judged by measuring the Fv/Fm ratio, decreased significantly at LTs higher than 38 °C. A post-illumination heat-induced reduction of the plastoquinone pool was observed in the 25 °C-plants, but not in the 35 °C-plants. Inhibition of Pn by heat stress correlated with a reduction of the activation state of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco). Western-blot analysis of Rubisco activase showed that heat stress resulted in a redistribution of activase polypeptides from the soluble to the insoluble fraction of extracts. Heat-dependent inhibition of Pn and PET could be reduced by increasing the intercellular CO2 concentration, but much more effectively so in the 35 °C-plants than in the 25 °C-plants. The 35 °C-plants recovered more efficiently from heat-dependent inhibition of Pn than the 25 °C-plants. The results show that growth at moderately high temperature hardly diminished inhibition of Pn by heat stress that originated from a reversible heat-dependent reduction of the Rubisco activation state. However, by improving the thermal stability of the thylakoid membranes it allowed the photosynthetic apparatus to preserve its functional potential at high LTs, thus minimizing the after-effects of heat stress.

Application and comparison of UTCI and PET in temperature climate conditions

The assessment of the thermal bioclimate is based on the human energy balance and derived indices such as Physiologically equivalent temperature (Pet) or Universal thermal Climate index (UtCi). These two indices were compared over a period often year based on hourly data in a middle european city with a temperate climate. The analysis performed shows that the differences obtained result from the different thermo-physiological settings of clothing insulation. For conditions with extremely high vapour pressure values, UtCi yields higher values than Pet, which could describe the thermo-physiological stress more appropriately.

Temperature and precipitation effects on δ13C depth profiles in SOM under temperate beech forests

Enrichment of 13C in SOM with soil depth is related to interacting processes influenced by temperature and precipitation. Our objectives were to derive climate effects on patterns of vertical δ13C values of soil organic matter (SOM) while minimizing the effect of confounding variables. We investigated vertical changes in δ13C values of SOM in 1-cm depth intervals in silvicultural mature beech (Fagus sylvatica L.) forest ecosystems in northern Rhineland-Palatinate across gradients of MAT (7.9 to 9.7 °C mean annual temperature) and MAP (607 to 1085 mm mean annual precipitation) in winter 2011. Forest stands (n = 10) were chosen based on data sets provided by the Rhineland-Palatinate Forest Administration so that variations in these gradients occurred while other environmental factors like physico-chemical soil properties, tree species, stand age, exposition and precipitation (for the temperature gradient) or temperature (for the precipitation gradient) did not differ among study sites. From litter down to the mineral soil at 10 cm depth, soil organic carbon (SOC) content decreased (47.5 ± SE 0.1% to 2.5 ± 0.1%) while the δ13C values increased (− 29.4 ± 0.1‰ to − 26.1 ± 0.1‰). Litter of sites under higher MAP/lower MAT had lower δ13C values which was in line with literature data on climate driven plant physiological process. To compare the dimension of the vertical 13C enrichment, δ13C values were regressed linearly against log-transformed carbon contents yielding absolute values of these slopes (beta). Beta values ranged between 0.6 and 4.5 (range of r from − 0.7 to − 1.0; p < 0.01). Due to an assumed decay continuum and similar variations of δ13C values in litter and in 10 cm depth, we conclude that effects on isotope composition in the Oi layer continue vertically and therefore, δ13C values in litter do not solely control beta values. Beta values decreased with increasing MAT (r = − 0.83; p < 0.05). Reduced soil moisture and therefore both, reduced microbial activity and reduced downward transport of microbial cycled DOM (=13C enriched) might be responsible for less pronounced δ13C depth profiles in case of high temperatures. Greater C:N ratios (lower degradability) of the litter under higher temperatures likely contributed to these depth trends. Beta values increased with increasing MAP (r = 0.73; p < 0.05). We found decreasing C:N ratios in the mineral soil that possibly indicates higher decomposition under higher precipitation. Exclusion of the organic layers from linear regressions indicated a stronger impact of MAP on the development of δ13C depth profiles. Our results confirm temperature and precipitation effects on δ13C depth profiles and indicate stronger 13C enrichment under lower MAT/higher MAP. Therefore, time series of vertical δ13C depth profiles might provide insights into climate change effects.