ALBERT

Description: Evergreen trees in the Mediterranean region must cope with a wide range of environmental stresses from summer drought to winter cold. The mildness of Mediterranean winters can periodically lead to favourable environmental conditions above the threshold for a positive carbon balance, benefitting evergreen woody species more than deciduous ones. The comparatively lower solar energy input in winter decreases the foliar light saturation point. This leads to a higher susceptibility to photoinhibitory stress especially when chilly (〈 12 °C) or freezing temperatures (〈 0 °C) coincide with clear skies and relatively high solar irradiances. Nonetheless, the advantage of evergreen species that are able to photosynthesize all year round where a significant fraction can be attributed to winter months, compensates for the lower carbon uptake during spring and summer in comparison to deciduous species. We investigated the ecophysiological behaviour of three co-occurring mature evergreen tree species (Quercus ilex L., Pinus halepensis Mill., and Arbutus unedo L.) during a period of mild winter conditions and their responses to a sudden cold period. The state of the photosynthetic machinery in both periods was thus tested by estimating the foliar photosynthetic potential with CO2 response curves in parallel with chlorophyll fluorescence measurements. The studied evergreen tree species benefited strongly from mild winter conditions by exhibiting extraordinarily high photosynthetic potentials similar to those under spring conditions. A sudden period of frost, however, negatively affected the photosynthetic apparatus, leading to significant decreases in key physiological parameters such as the maximum carboxylation velocity (Vc, max), the maximum photosynthetic electron transport rate (Jmax), and the optimal fluorometric quantum yield of photosystem II (Fv/Fm). This change persisted for several weeks after the cold period despite the recovery of the temperature to the conditions previous to the frost event. The responses of Vc, max and Jmax were highly species-specific, where Q. ilex exhibited the highest and P. halepensis the lowest reductions. In contrast, the optimal fluorometric quantum yield of photosystem II (Fv/Fm) was significantly lower in A. unedo after the cold period. The leaf position played an important role in Q. ilex showing a comparatively stronger winter effect on sunlit leaves. Our results generally agreed with the previous classifications of photoinhibition-avoiding (P. halepensis) and photoinhibition-tolerant (Q. ilex) species on the basis of their susceptibility to dynamic photoinhibition, whereas A. unedo was the least tolerant to photoinhibition, which was chronic in this species. Q. ilex and P. halepensis seem to follow contrasting photoprotective strategies which are, however, equally successful under the prevailing conditions exhibiting an adaptive advantage over A. unedo in our study site. These results show that our understanding of the dynamics of interspecific competition in Mediterranean ecosystems requires consideration of the physiological behaviour during winter which may have important implications for long-term carbon budgets and growth trends.

Description: Tropical cyclones (typhoons/hurricanes) have major impacts on the biogeochemistry of forest ecosystems, but the stochastic nature and the long intervals between storms means that there are limited data on their effects. We characterised the impacts of 14 typhoons over six years on hydrochemistry of a subtropical forest plantation in Taiwan, a region experiencing frequent typhoons. Typhoons contributed 1/3 of the annual rainfall on average, but ranged from 4 to 55%. The stochastic nature of annual typhoon related precipitation poses a challenge with respect to managing the impacts of these extreme events. This challenge is exacerbated by the fact that typhoon-related rainfall is not significantly correlated with wind velocity, the current focus of weather forecasts. Thus, little advance warning is provided for the hydrological impacts of these storms. The typhoons we studied contributed approximately one third of the annual input and output of most nutrients (except nitrogen) during an average 9.5 day yr−1 period, resulting in nutrient input/output rates an order of magnitude greater than during non-typhoon months. Nitrate output balanced input during the non-typhoon period, but during the typhoon period an average of 10 kg ha−1 yr−1 nitrate was lost. Streamwater chemistry exhibited similarly high variability during typhoon and non-typhoon periods and returned to pre-typhoon levels one to three weeks following each typhoon. The streamwater chemistry appears to be very resilient in response to typhoons, resulting in minimal loss of nutrients.

Description: Tropical cyclones (typhoons/hurricanes) have major impacts on the biogeochemistry of forest ecosystems, but the stochastic nature and the long intervals between storms means that there are limited data on their effects. We characterized the impacts of 14 typhoons over six years on hydrochemistry of a subtropical forest plantation in Taiwan, a region experiencing frequent typhoons. Typhoons contributed 1/3 of annual rainfall on average, but ranged from 4% to 55%. The stochastic nature of annual typhoon related precipitation poses a challenge with respect to managing the impacts of these extreme events. This challenge is exacerbated by the fact that typhoon-related rainfall is not significantly correlated with wind velocity, the current focus of weather forecasts. Thus little advance warning is provided for the hydrological impacts of these storms. The typhoons we studied contributed approximately one third of the annual input and output of most nutrients (except nitrogen) during an average 9.5d yr−1 period, resulting in nutrient input/output rates an order of magnitude greater than during non-typhoon period. Nitrate output balanced input during the non-typhoon period, but during the typhoon period an average of 10 kg ha−1 yr−1 nitrate was lost. Streamwater chemistry exhibited similarly high variability during typhoon and non-typhoon periods and returned to pre-typhoon levels one to three weeks following each typhoon. The streamwater chemistry appears to be very resilient in response to typhoons, resulting in minimal loss of nutrients.

Description: Soil respiration (SR) is a major component of ecosystem's carbon cycle and represents the second largest CO2 flux of the terrestrial biosphere. Soil temperature is considered to be the primary control on SR whereas soil moisture as the secondary control factor. However, soil moisture can become the dominant control on SR in very wet or dry conditions. Determining the trigger that switches-on soil moisture as the primary control factor of SR will provide a deeper understanding on how SR changes under projected future increased droughts. Specific objectives of this study were (1) to investigate the seasonal variations and the relationship between SR and both soil temperature and moisture in a Mediterranean riparian forest along a groundwater level gradient; (2) to determine soil moisture thresholds at which SR is rather controlled by soil moisture than by temperature; (3) to compare SR responses under different tree species present in a Mediterranean riparian forest (Alnus, glutinosa, Populus nigra and Fraxinus excelsior). Results showed that the heterotrophic soil respiration rate, groundwater level and 30 cm integral soil moisture (SM30) decreased significantly from riverside to uphill and showed a pronounced seasonality. SR rates showed significant differences among tree species, with higher SR for P. nigra and lower SR for A. glutinosa. The lower threshold of soil moisture was 20 and 17% for heterotrophic and total SR respectively. Daily mean SR rate was positively correlated with soil temperature when soil moisture exceeded the threshold, with Q10 values ranging from 1.19 to 2.14; nevertheless, SR became decoupled from soil temperature when soil moisture dropped below these thresholds.

Description: Soil respiration (SR) is a major component of ecosystems' carbon cycles and represents the second largest CO2 flux in the terrestrial biosphere. Soil temperature is considered to be the primary abiotic control on SR, whereas soil moisture is the secondary control factor. However, soil moisture can become the dominant control on SR in very wet or dry conditions. Determining the trigger that makes soil moisture as the primary control factor of SR will provide a deeper understanding on how SR changes under the projected future increase in droughts. Specific objectives of this study were (1) to investigate the seasonal variations and the relationship between SR and both soil temperature and moisture in a Mediterranean riparian forest along a groundwater level gradient; (2) to determine soil moisture thresholds at which SR is controlled by soil moisture rather than by temperature; (3) to compare SR responses under different tree species present in a Mediterranean riparian forest (Alnus glutinosa, Populus nigra and Fraxinus excelsior). Results showed that the heterotrophic soil respiration rate, groundwater level and 30 cm integral soil moisture (SM30) decreased significantly from the riverside moving uphill and showed a pronounced seasonality. SR rates showed significant differences between tree species, with higher SR for P. nigra and lower SR for A. glutinosa. The lower threshold of soil moisture was 20 and 17% for heterotrophic and total SR, respectively. Daily mean SR rate was positively correlated with soil temperature when soil moisture exceeded the threshold, with Q10 values ranging from 1.19 to 2.14; nevertheless, SR became decoupled from soil temperature when soil moisture dropped below these thresholds.

Description: Evergreen trees in the Mediterranean region must cope with a wide range of environmental stresses from summer drought to winter cold. The mildness of Mediterranean winters can periodically lead to favourable environmental conditions above the threshold for a positive carbon balance, benefitting evergreen woody species more than deciduous ones. The comparatively lower solar energy input in winter decreases the foliar light saturation point. This leads to a higher susceptibility to photoinhibitory stress especially when chilly (〈 12 °C) or freezing temperatures (〈 0 °C) coincide with clear skies and relatively high solar irradiances. Nonetheless, the advantage of evergreen species that are able to photosynthesize all year round where a significant fraction can be attributed to winter months, compensates for the lower carbon uptake during spring and summer in comparison to deciduous species. We investigated the ecophysiological behaviour of three co-occurring mature evergreen tree species (Quercus ilex L., Pinus halepensis Mill., and Arbutus unedo L.). Therefore, we collected twigs from the field during a period of mild winter conditions and after a sudden cold period. After both periods, the state of the photosynthetic machinery was tested in the laboratory by estimating the foliar photosynthetic potential with CO2 response curves in parallel with chlorophyll fluorescence measurements. The studied evergreen tree species benefited strongly from mild winter conditions by exhibiting extraordinarily high photosynthetic potentials. A sudden period of frost, however, negatively affected the photosynthetic apparatus, leading to significant decreases in key physiological parameters such as the maximum carboxylation velocity (Vc, max), the maximum photosynthetic electron transport rate (Jmax), and the optimal fluorometric quantum yield of photosystem II (Fv/Fm). The responses of Vc, max and Jmax were highly species specific, with Q. ilex exhibiting the highest and P. halepensis the lowest reductions. In contrast, the optimal fluorometric quantum yield of photosystem II (Fv/Fm) was significantly lower in A. unedo after the cold period. The leaf position played an important role in Q. ilex showing a stronger winter effect on sunlit leaves in comparison to shaded leaves. Our results generally agreed with the previous classifications of photoinhibition-tolerant (P. halepensis) and photoinhibition-avoiding (Q. ilex) species on the basis of their susceptibility to dynamic photoinhibition, whereas A. unedo was the least tolerant to photoinhibition, which was chronic in this species. Q. ilex and P. halepensis seem to follow contrasting photoprotective strategies. However, they seemed equally successful under the prevailing conditions exhibiting an adaptive advantage over A. unedo. These results show that our understanding of the dynamics of interspecific competition in Mediterranean ecosystems requires consideration of the physiological behaviour during winter which may have important implications for long-term carbon budgets and growth trends.