It is not just a 'trade-off': indications for sink- and source-limitation to vegetative and regenerative growth in an old-growth beech forest.

Controls on tree growth are key issues in plant physiology. The hypothesis of our study was that the interannual variability of wood and fruit production are primarily controlled directly by weather conditions (sink limitation), while carbon assimilation (source limitation) plays a secondary role. We analyzed the interannual variability of weather conditions, gross primary productivity (GPP) and net primary productivity (NPP) of wood and fruits of an old-growth, unmanaged Fagus sylvatica forest over 14 yr, including six mast years. In a multiple linear regression model, c. 71% of the annual variation in wood-NPP could be explained by mean air temperature in May, precipitation from April to May (positive influence) and fruit-NPP (negative influence). GPP of June to July solely explained c. 42% of the variation in wood-NPP. Fruit-NPP was positively related to summer precipitation 2 yr before (R2  = 0.85), and negatively to precipitation in May (R2  = 0.83) in the fruit years. GPP had no influence on fruit-NPP. Our results suggest a complex system of sink and source limitations to tree growth driven by weather conditions and going beyond a simple carbon-mediated 'trade-off' between regenerative and vegetative growth.

An empirical study of the wound effect on sap flux density measured with thermal dissipation probes.

The insertion of thermal dissipation (TD) sensors on tree stems for sap flux density (SFD) measurements can lead to SFD underestimations due to a wound formation close to the drill hole. However, the wound effect has not been assessed experimentally for this method yet. Here, we propose an empirical approach to investigate the effect of the wound healing on measured sap flux with TD probes. The approach was performed for both, diffuse-porous (Fagus sylvatica (Linnaeus)) and ring-porous (Quercus petraea (Lieblein)) species. Thermal dissipation probes were installed on different dates along the growing season to document the effects of the dynamic wound formation. The trees were cut in autumn and additional sensors were installed in the cut stems, therefore, without potential effects of wound development. A range of water pressures was applied to the stem segments and SFDs were simultaneously measured by TD sensors as well as gravimetrically in the laboratory. The formation of wounds around sensors installed in living tree stems led to underestimation of SFD by 21.4 ± 3 and 47.5 ± 3.8% in beech and oak, respectively. The differences between SFD underestimations of diffuse-porous beech and ring-porous oak were, however, not statistically significant. Sensors with 5-, 11- and 22-week-old wounds also showed no significant differences, which implies that the influence of wound formation on SFD estimates was completed within the first few weeks after perforation. These results were confirmed by time courses of SFD measurements in the field. Field SFD values decreased immediately after sensor installation and reached stable values after ~2 weeks with similar underestimations to the ones observed in the laboratory. We therefore propose a feasible approach to correct directly field observations of SFD for potential underestimations due to the wound effect.

Remotely-sensed detection of effects of extreme droughts on gross primary production.

Severe droughts strongly impact photosynthesis (GPP), and satellite imagery has yet to demonstrate its ability to detect drought effects. Especially changes in vegetation functioning when vegetation state remains unaltered (no browning or defoliation) pose a challenge to satellite-derived indicators. We evaluated the performance of different satellite indicators to detect strong drought effects on GPP in a beech forest in France (Hesse), where vegetation state remained largely unaffected while GPP decreased substantially. We compared the results with three additional sites: a Mediterranean holm oak forest (Puéchabon), a temperate beech forest (Hainich), and a semi-arid grassland (Bugacpuszta). In Hesse, a three-year reduction in GPP following drought was detected only by the Enhanced Vegetation Index (EVI). The Photochemical Reflectance Index (PRI) also detected this drought effect, but only after normalization for absorbed light. In Puéchabon normalized PRI outperformed the other indicators, while the short-term drought effect in Hainich was not detected by any tested indicator. In contrast, most indicators, but not PRI, captured the drought effects in Bugacpuszta. Hence, PRI improved detection of drought effects on GPP in forests and we propose that PRI normalized for absorbed light is considered in future algorithms to estimate GPP from space.

Potential and limitations of inferring ecosystem photosynthetic capacity from leaf functional traits.

The aim of this study was to systematically analyze the potential and limitations of using plant functional trait observations from global databases versus in situ data to improve our understanding of vegetation impacts on ecosystem functional properties (EFPs). Using ecosystem photosynthetic capacity as an example, we first provide an objective approach to derive robust EFP estimates from gross primary productivity (GPP) obtained from eddy covariance flux measurements. Second, we investigate the impact of synchronizing EFPs and plant functional traits in time and space to evaluate their relationships, and the extent to which we can benefit from global plant trait databases to explain the variability of ecosystem photosynthetic capacity. Finally, we identify a set of plant functional traits controlling ecosystem photosynthetic capacity at selected sites. Suitable estimates of the ecosystem photosynthetic capacity can be derived from light response curve of GPP responding to radiation (photosynthetically active radiation or absorbed photosynthetically active radiation). Although the effect of climate is minimized in these calculations, the estimates indicate substantial interannual variation of the photosynthetic capacity, even after removing site-years with confounding factors like disturbance such as fire events. The relationships between foliar nitrogen concentration and ecosystem photosynthetic capacity are tighter when both of the measurements are synchronized in space and time. When using multiple plant traits simultaneously as predictors for ecosystem photosynthetic capacity variation, the combination of leaf carbon to nitrogen ratio with leaf phosphorus content explains the variance of ecosystem photosynthetic capacity best (adjusted R2 = 0.55). Overall, this study provides an objective approach to identify links between leaf level traits and canopy level processes and highlights the relevance of the dynamic nature of ecosystems. Synchronizing measurements of eddy covariance fluxes and plant traits in time and space is shown to be highly relevant to better understand the importance of intra- and interspecific trait variation on ecosystem functioning.

Estimating daytime ecosystem respiration from eddy-flux data.

To understand what governs the patterns of net ecosystem exchange of CO2, an understanding of factors influencing the component fluxes, ecosystem respiration and gross primary production is needed. In the present paper, we introduce an alternative method for estimating daytime ecosystem respiration based on whole ecosystem fluxes from a linear regression of photosynthetic photon flux density data vs. daytime net ecosystem exchange data at forest ecosystem level. This method is based on the principles of the Kok-method applied at leaf level for estimating daytime respiration. We demonstrate the method with field data and provide a discussion of the limitations of the method.

Comparative measurements of transpiration and canopy conductance in two mixed deciduous woodlands differing in structure and species composition.

Transpiration of two heterogeneous broad-leaved woodlands in southern England was monitored by the sap flux technique throughout the 2006 growing season. Grimsbury Wood, which had a leaf area index (LAI) of 3.9, was dominated by oak (Quercus robur L.) and birch (Betula pubescens L.) and had a continuous hazel (Corylus avellana L.) understory. Wytham Woods, which had an LAI of 3.6, was dominated by ash (Fraxinus excelsior L.) and sycamore (Acer pseudoplatanus L.) and had only a sparse understory. Annual canopy transpiration was 367 mm for Grimsbury Wood and 397 mm for Wytham Woods. These values were similar to those for beech (Fagus sylvatica L.) plantations in the same region, and differ from one another by less than the typical margin of uncertainty of the sap flux technique. Canopy conductance (g(c)), calculated for both woodlands by inverting the Penman-Monteith equation, was related to incoming solar radiation (R(G)) and the vapor pressure deficit (D). The response of g(c) to R(G) was similar for both forests. Both reference conductance (g(cref)), defined as g(c) at D=1 kPa, and stomatal sensitivity (-m), defined as the slope of the logarithmic response curve of g(c) to D, increased during the growing season at Wytham Woods but not at Grimsbury Wood. The -m/g(cref) ratio was significantly lower at Wytham Woods than at Grimsbury Wood and was insufficient to keep the difference between leaf and soil water potentials constant, according to a simple hydraulic model. This meant that annual water consumption of the two woodlands was similar despite different regulatory mechanisms and associated short-term variations in canopy transpiration. The -m/g(cref) ratio depended on the range of D under which the measurements were made. This was shown to be particularly important for studies conducted under low and narrow ranges of D.

Seasonal and interannual variability of canopy transpiration of a hedgerow in southern England.

Transpiration from a hawthorn (Crataegus monogyna L.) dominated hedgerow in southern England was measured continuously over two growing seasons by the sap flow technique. Accompanying measurements of structural parameters, microclimate and leaf stomatal and boundary layer conductances were used to establish the driving factors of hedgerow transpiration. Observed transpiration rates, reaching peak values of around 8 mm day(-1) and a seasonal mean of about 3.5 mm day(-1), were higher than those reported for most other temperate deciduous woodlands, except short-rotation coppice and wet woodlands. The high rates were caused by the structural and physiological characteristics of hawthorn leaves, which exhibited much higher stomatal and boundary-layer conductances than those of the second-most abundant woody species in the hedgerow, field maple (Acer campestre L.). Only in the hot summer of 2003 did stomatal conductance, and thus transpiration, decrease substantially. The hedgerow canopy was always closely coupled to the atmosphere. Hedgerow transpiration equaled potential evaporation (calculated by the Priestley-Taylor formula) in 2003 and exceeded it in 2004, which meant that a substantial fraction of the energy (21% in 2003 and more than 37% in 2004) came from advection. Hedgerow canopy conductance (g(c)), as inferred from the sap flow data by inverting the Penman-Monteith equation, responded to solar radiation (R(G)) and vapor pressure deficit (D). Although the response to R(G) showed no systematic temporal variation, the response to D, described as g(c)(D) = g(cref) - mln(D), changed seasonally. The reference g(c) depended on leaf area index and the ratio of -m/g(cref) on long-term mean daytime D. A model is proposed based on these observations that predicts canopy conductance for the hawthorn hedge from standard weather data.