Differentiating drought legacy effects on vegetation growth over the temperate Northern Hemisphere.

In view of future changes in climate, it is important to better understand how different plant functional groups (PFGs) respond to warmer and drier conditions, particularly in temperate regions where an increase in both the frequency and severity of drought is expected. The patterns and mechanisms of immediate and delayed impacts of extreme drought on vegetation growth remain poorly quantified. Using satellite measurements of vegetation greenness, in-situ tree-ring records, eddy-covariance CO2 and water flux measurements, and meta-analyses of source water of plant use among PFGs, we show that drought legacy effects on vegetation growth differ markedly between forests, shrubs and grass across diverse bioclimatic conditions over the temperate Northern Hemisphere. Deep-rooted forests exhibit a drought legacy response with reduced growth during up to 4 years after an extreme drought, whereas shrubs and grass have drought legacy effects of approximately 2 years and 1 year, respectively. Statistical analyses partly attribute the differences in drought legacy effects among PFGs to plant eco-hydrological properties (related to traits), including plant water use and hydraulic responses. These results can be used to improve the representation of drought response of different PFGs in land surface models, and assess their biogeochemical and biophysical feedbacks in response to a warmer and drier climate.

Disaggregation of canopy photosynthesis among tree species in a mixed broadleaf forest.

Carbon dioxide sequestration from the atmosphere is commonly assessed using the eddy covariance method. Its net flux signal can be decomposed into gross primary production and ecosystem respiration components, but these have seldom been tested against independent methods. In addition, eddy covariance lacks the ability to partition carbon sequestration among individual trees or species within mixed forests. Therefore, we compared gross primary production from eddy covariance versus an independent method based on sap flow and water-use efficiency, as measured by the tissue heat balance method and δ13C of phloem contents, respectively. The latter measurements were conducted on individual trees throughout a growing season in a mixed broadleaf forest dominated by three tree species, namely English oak, narrow-leaved ash and common hornbeam (Quercus robur L., Fraxinus angustifolia Vahl, and Carpinus betulus L., respectively). In this context, we applied an alternative ecophysiological method aimed at verifying the accuracy of a state-of-the-art eddy covariance system while also offering a solution to the partitioning problem. We observed strong agreement in the ecosystem gross primary production estimates (R2 = 0.56; P < 0.0001), with correlation being especially high and nearly on the 1:1 line in the period before the end of July (R2 = 0.85; P < 0.0001). After this period, the estimates of gross primary production began to diverge. Possible reasons for the divergence are discussed, focusing especially on phenology and the limitation of the isotopic data. English oak showed the highest per-tree daily photosynthetic rates among tree species, but the smaller, more abundant common hornbeam contributed most to the stand-level summation, especially early in the spring. These findings provide a rigorous test of the methods and the species-level photosynthesis offers avenues for enhancing forest management aimed at carbon sequestration.

Seasonally varying relationship between stem respiration, increment and carbon allocation of Norway spruce trees.

Stem respiration is an important component of an ecosystem's carbon budget. Beside environmental factors, it depends highly on tree energy demands for stem growth. Determination of the relationship between stem growth and stem respiration would help to reveal the response of stem respiration to changing climate, which is expected to substantially affect tree growth. Common measurement of stem radial increment does not record all aspects of stem growth processes, especially those connected with cell wall thickening; therefore, the relationship between stem respiration and stem radial increment may vary depending on the wood cell growth differentiation phase. This study presents results from measurements of stem respiration and increment carried out for seven growing seasons in a young Norway spruce forest. Moreover, rates of carbon allocation to stems were modeled for these years. Stem respiration was divided into maintenance (Rm) and growth respiration (Rg) based upon the mature tissue method. There was a close relationship between Rg and daily stem radial increment (dSRI), and this relationship differed before and after dSRI seasonal maximum, which was around 19 June. Before this date, Rg increased exponentially with dSRI, while after this date logarithmically. This is a result of later maxima of Rg and its slower decrease when compared with dSRI, which is connected with energy demands for cell wall thickening. Rg reached a maxima at the end of June or in July. The maximum of carbon allocation to stem peaked in late summer, when Rg mostly tended to decrease. The overall contribution of Rg to stem CO2 efflux amounted to 46.9% for the growing period from May to September and 38.2% for the year as a whole. This study shows that further deeper analysis of in situ stem growth and stem respiration dynamics is greatly needed, especially with a focus on wood formation on a cell level.

Analysis of floodplain forest sensitivity to drought.

Floodplain forests are very complex, productive ecosystems, capable of storing huge amounts of soil carbon. With the increasing occurrence of extreme events, they are today among the most threatened ecosystems. Our study's main goal was to assess the productivity of a floodplain forest located at Lanzhot in the Czech Republic from two perspectives: carbon uptake (using an eddy covariance method) and stem radius variations (using dendrometers). We aimed to determine which conditions allow for high ecosystem production and what role drought plays in reducing such production potential. Additionally, we were interested to determine the relative soil water content threshold indicating the onset and duration of this event. We hypothesized that summer drought in 2018 had the most significant negative effects on the overall annual carbon and water budgets. In contrast with our original hypothesis, we found that an exceptionally warm spring in 2018 caused a positive gross primary production (GPP) and evapotranspiration (ET) anomaly that consequently led in 2018 to the highest seasonal total GPP and ET from all of the investigated years (2015-2018). The results showed ring-porous species to be the most drought resistant. Relative soil water content threshold of approximately 0.45 was determined as indicating the onset of drought stress. This article is part of the theme issue 'Impacts of the 2018 severe drought and heatwave in Europe: from site to continental scale'.

Variability of stem CO2 efflux response to temperature over the diel period.

This study presents results from continuous measurements of stem CO2 efflux carried out for seven growing seasons in a young Norway spruce forest. The objective of the study was to determine differences in temperature sensitivity of stem CO2 efflux (Q10) during night (when sap flow is zero or nearly zero), during early afternoon (when the maximum rate of sap flow occurs) and during two transition periods between the aforementioned periods. The highest Q10 was recorded during the period of zero sap flow, while the lowest Q10 was observed in period of the highest sap flow. Calculating Q10 using only data from the period of zero sap flow resulted in a Q10 that was higher by as much as 19% compared with Q10 calculated using 24 h data. On the other hand, basing the calculation on data from the period of the highest sap flow yielded 5.6% lower Q10 than if 24 h data were used. Considering that change in CO2 efflux lagged in time behind changing stem temperature, there was only a small effect on calculated Q10 for periods with zero and the highest sap flow. A larger effect of the time lag (by as much as 15%) was observed for the two transition periods. Stem CO2 efflux was modelled based on the night CO2 efflux response to temperature. This model had a tendency to overestimate CO2 efflux during daytime, thus indicating potential daytime depression of stem CO2 efflux compared with the values predicated on the basis of temperature caused by CO2 transport upward in the sap flow. This view was supported by our results inasmuch as the overestimation grew with sap flow that was modelled on the basis of photosynthetically active radiation and vapour pressure deficit.

Seasonal variation in CO2 efflux of stems and branches of Norway spruce trees.

BACKGROUND AND AIMS: Stem and branch respiration, important components of total forest ecosystem respiration, were measured on Norway spruce (Picea abies) trees from May to October in four consecutive years in order (1) to evaluate the influence of temperature on woody tissue CO2 efflux with special focus on variation in Q10 (change in respiration rate resulting from a 10 degrees C increase in temperature) within and between seasons, and (2) to quantify the contribution of above-ground woody tissue (stem and branch) respiration to the carbon balance of the forest ecosystem. METHODS: Stem and branch CO2 efflux were measured, using an IRGA and a closed gas exchange system, 3-4 times per month on 22-year-old trees under natural conditions. Measurements of ecosystem CO2 fluxes were also determined during the whole experiment by using the eddy covariance system. Stem and branch temperatures were monitored at 10-min intervals during the whole experiment. KEY RESULTS: The temperature of the woody tissue of stems and branches explained up to 68% of their CO2 efflux. The mean annual Q10 values ranged from 2.20 to 2.32 for stems and from 2.03 to 2.25 for branches. The mean annual normalized respiration rate, R10, for stems and branches ranged from 1.71 to 2.12 micromol CO2 m(-2)s (-1) and from 0.24 to 0.31 micromol CO2 m(-2) s(-1), respectively. The annual contribution of stem and branch CO2 efflux to total ecosystem respiration were, respectively, 8.9 and 8.1% in 1999, 9.2 and 9.2% in 2000, 7.6 and 8.6% in 2001, and 8.6 and 7.9% in 2002. Standard deviation for both components ranged from 3 to 8% of the mean. CONCLUSIONS: Stem and branch CO2 efflux varied diurnally and seasonally, and were related to the temperature of the woody tissue and to growth. The proportion of CO2 efflux from stems and branches is a significant component of the total forest ecosystem respiration, approx. 8% over the 4 years, and predictive models must take their contribution into account.

Variability in temperature dependence of stem CO2 efflux from Norway spruce trees.

This study presents results from continuous measurements of stem CO2 efflux carried out for seven experimental seasons (from May to October) in a young Norway spruce forest. The objectives of the study were to determine variability in the response of stem CO2 efflux to stem temperature over the season and to observe differences in the calculated relationship between stem temperature and CO2 efflux based on full growing season data or on data divided into periods according to stem growth rate. Temperature sensitivity of stem CO2 efflux (Q10) calculated for the established periods ranged between 1.61 and 3.46 and varied over the season, with the lowest values occurring in July and August. Q10 calculated using data from the full growing seasons ranged between 2.30 and 2.94 and was often significantly higher than Q10 calculated for the individual periods. Temperature-normalized stem CO2 efflux (R10) determined using Q10 from growing season data was overestimated when the temperature was below 10 °C and underestimated when the temperature was above 10 °C, compared with R10 calculated using Q10 established for the individual periods. The differences in daily mean R10 calculated by these two approaches ranged between -0.9 and 0.2 µmol CO2 m-2 s-1. The results of this study confirm that long periods for determining the temperature dependence of stem CO2 efflux encompass different statuses of the wood (especially in relation to stem growth). This may cause bias in models using this relationship for estimating stem CO2 efflux as a function of temperature.

Extreme warm temperatures alter forest phenology and productivity in Europe.

Recent climate warming has shifted the timing of spring and autumn vegetation phenological events in the temperate and boreal forest ecosystems of Europe. In many areas spring phenological events start earlier and autumn events switch between earlier and later onset. Consequently, the length of growing season in mid and high latitudes of European forest is extended. However, the lagged effects (i.e. the impact of a warm spring or autumn on the subsequent phenological events) on vegetation phenology and productivity are less explored. In this study, we have (1) characterised extreme warm spring and extreme warm autumn events in Europe during 2003-2011, and (2) investigated if direct impact on forest phenology and productivity due to a specific warm event translated to a lagged effect in subsequent phenological events. We found that warmer events in spring occurred extensively in high latitude Europe producing a significant earlier onset of greening (OG) in broadleaf deciduous forest (BLDF) and mixed forest (MF). However, this earlier OG did not show any significant lagged effects on autumnal senescence. Needleleaf evergreen forest (NLEF), BLDF and MF showed a significantly delayed end of senescence (EOS) as a result of extreme warm autumn events; and in the following year's spring phenological events, OG started significantly earlier. Extreme warm spring events directly led to significant (p=0.0189) increases in the productivity of BLDF. In order to have a complete understanding of ecosystems response to warm temperature during key phenological events, particularly autumn events, the lagged effect on the next growing season should be considered.

Carbon exchange between ecosystems and atmosphere in the Czech Republic is affected by climate factors.

By comparing five ecosystem types in the Czech Republic over several years, we recorded the highest carbon sequestration potential in an evergreen Norway spruce forest (100%) and an agroecosystem (65%), followed by European beech forest (25%) and a wetland ecosystem (20%). Because of a massive ecosystem respiration, the final carbon gain of the grassland was negative. Climate was shown to be an important factor of carbon uptake by ecosystems: by varying the growing season length (a 22-d longer season in 2005 than in 2007 increased carbon sink by 13%) or by the effect of short- term synoptic situations (e.g. summer hot and dry days reduced net carbon storage by 58% relative to hot and wet days). Carbon uptake is strongly affected by the ontogeny and a production strategy which is demonstrated by the comparison of seasonal course of carbon uptake between coniferous (Norway spruce) and deciduous (European beech) stands.