Summary• Phloem is the main pathway for transferring photosynthates belowground. In situ 13 C pulse labelling of trees 8-10 m tall was conducted in the field on 10 beech (Fagus sylvatica) trees, six sessile oak (Quercus petraea) trees and 10 maritime pine (Pinus pinaster) trees throughout the growing season.• Respired 13 CO 2 from trunks was tracked at different heights using tunable diode laser absorption spectrometry to determine time lags and the velocity of carbon transfer (V). The isotope composition of phloem extracts was measured on several occasions after labelling and used to estimate the rate constant of phloem sap outflux (k P ).• Pulse labelling together with high-frequency measurement of the isotope composition of trunk CO 2 efflux is a promising tool for studying phloem transport in the field. Seasonal variability in V was predicted in pine and oak by bivariate linear regressions with air temperature and soil water content. V differed among the three species consistently with known differences in phloem anatomy between broadleaf and coniferous trees.• V increased with tree diameter in oak and beech, reflecting a nonlinear increase in volumetric flow with increasing bark cross-sectional area, which suggests changes in allocation pattern with tree diameter in broadleaf species. Discrepancies between V and k P indicate vertical changes in functional phloem properties.
Drought and heat events, such as the 2018 European drought, interact with the exchange of energy between the land surface and the atmosphere, potentially affecting albedo, sensible and latent heat fluxes, as well as CO 2 exchange. Each of these quantities may aggravate or mitigate the drought, heat, their side effects on productivity, water scarcity and global warming. We used measurements of 56 eddy covariance sites across Europe to examine the response of fluxes to extreme drought prevailing most of the year 2018 and how the response differed across various ecosystem types (forests, grasslands, croplands and peatlands). Each component of the surface radiation and energy balance observed in 2018 was compared to available data per site during a reference period 2004–2017. Based on anomalies in precipitation and reference evapotranspiration, we classified 46 sites as drought affected. These received on average 9% more solar radiation and released 32% more sensible heat to the atmosphere compared to the mean of the reference period. In general, drought decreased net CO 2 uptake by 17.8%, but did not significantly change net evapotranspiration. The response of these fluxes differed characteristically between ecosystems; in particular, the general increase in the evaporative index was strongest in peatlands and weakest in croplands. This article is part of the theme issue ‘Impacts of the 2018 severe drought and heatwave in Europe: from site to continental scale’.
Using nurse shrubs to improve tree seedling establishment in stressed environments is a common practice in forestry. Recent refinements of the stress‐gradient hypothesis suggest that positive nurse effects occur under intermediate stress and decline in the harshest conditions. Additionally, indirect facilitation (e.g. protection from herbivory) is expected in low‐stressed/productive systems according to the initial stress‐gradient hypothesis. However, there have been fewer investigations into the use of nurse shrubs to decrease herbivore impacts in stressed systems compared with the role of nurses in stress alleviation. This raises the need to clarify conditions in which the use of nurse shrubs would be most appropriate. We conducted a field experiment in the sand dune forest of southwest France characterized by severe summer water shortages and the presence of different types of wild herbivores. In 2015 and 2016, Pinus pinaster seedlings were planted under shrubs Arbutus unedo or in shrub‐removed locations, in fenced/unfenced areas. Survival, cause of mortality and water stress were monitored for 1 year after plantation. Summer drought was the main cause of mortality. Water stress increased in both years as summer progressed. We found direct facilitation only in July 2016 at intermediate water stress, due to a decrease in vapour pressure deficit under shrubs. These positive effects declined in late summer when stress was at a maximum. Rodent damage occurred under shrubs in spring whereas ungulate browsing was observed in open areas in autumn/winter. Overall seedling survival was equivalent under or outside shrubs during the first year, but was better under shrubs in the second one. Synthesis and applications. Our results partially validate the refinement of the stress‐gradient hypothesis, with a decline of positive interactions in the most water‐stressed conditions. This draws attention to the difficulty of using shrubs as nurses to improve tree regeneration in the most xeric systems. In addition, we show that indirect associational effects, both positive and negative, could be as important as abiotic stress alleviation in a stressful ecosystem, and using shrubs to facilitate seedling survival could be efficient depending on the type of herbivore involved. This highlights the fact that considering plant–plant interactions can help in the design of management options, providing that up‐to‐date ecological theories are considered and that the different biotic and abiotic constraints are accurately evaluated.
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