Integrating environmental gradients into breeding: application of genomic reactions norms in a perennial species.

Global warming threatens the productivity of forest plantations. We propose here the integration of environmental information into a genomic evaluation scheme using individual reaction norms, to enable the quantification of resilience in forest tree improvement and conservation strategies in the coming decades. Random regression models were used to fit wood ring series, reflecting the longitudinal phenotypic plasticity of tree growth, according to various environmental gradients. The predictive ability of the models was considered to select the most relevant environmental gradient, namely a gradient derived from an ecophysiological model and combining trunk water potential and temperature. Even if the individual ranking was preserved over most of the environmental gradient, strong genotype x environment interactions were detected in the extreme unfavorable part of the gradient, which includes environmental conditions that are very likely to be more frequent in the future. Combining genomic information and longitudinal data allowed to predict the growth of individuals in environments where they have not been observed. Phenotyping of 50% of the individuals in all the environments studied allowed to predict the growth of the remaining 50% of individuals in all these environments with a predictive ability of 0.25. Without changing the total number of observations, adding observations in a reduced number of environments for the individuals to be predicted, while decreasing the number of individuals phenotyped in all environments, increased the predictive ability to 0.59, highlighting the importance of phenotypic data allocation. We found that genomic reaction norms are useful for the characterization and prediction of the function of genetic parameters and facilitate breeding in a climate change context.

In situ assessment of the velocity of carbon transfer by tracing 13 C in trunk CO2 efflux after pulse labelling: variations among tree species and seasons.

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) CO2 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 (kP ). Pulse labelling together with high-frequency measurement of the isotope composition of trunk CO2 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 kP indicate vertical changes in functional phloem properties.

Photosynthetic carbon isotope discrimination and its relationship to the carbon isotope signals of stem, soil and ecosystem respiration.

• Photosynthetic carbon (C) isotope discrimination (Δ(Α)) labels photosynthates (δ(A) ) and atmospheric CO(2) (δ(a)) with variable C isotope compositions during fluctuating environmental conditions. In this context, the C isotope composition of respired CO(2) within ecosystems is often hypothesized to vary temporally with Δ(Α). • We investigated the relationship between Δ(Α) and the C isotope signals from stem (δ(W)), soil (δ(S)) and ecosystem (δ(E)) respired CO(2) to environmental fluctuations, using novel tuneable diode laser absorption spectrometer instrumentation in a mature maritime pine forest. • Broad seasonal changes in Δ(Α) were reflected in δ(W,) δ(S) and δ(E). However, respired CO(2) signals had smaller short-term variations than Δ(A) and were offset and delayed by 2-10 d, indicating fractionation and isotopic mixing in a large C pool. Variations in δ(S) did not follow Δ(A) at all times, especially during rainy periods and when there is a strong demand for C allocation above ground. • It is likely that future isotope-enabled vegetation models will need to develop transfer functions that can account for these phenomena in order to interpret and predict the isotopic impact of biosphere gas exchange on the C isotope composition of atmospheric CO(2).

Modeling climate change effects on the potential production of French plains forests at the sub-regional level.

We modeled the effects of climate change and two forest management scenarios on wood production and forest carbon balance in French forests using process-based models of forest growth. We combined data from the national forest inventory and soil network survey, which were aggregated over a 50 x 50-km grid, i.e., the spatial resolution of the climate scenario data. We predicted and analyzed the climate impact on potential forest production over the period 1960-2100. All models predicted a slight increase in potential forest yield until 2030-2050, followed by a plateau or a decline around 2070-2100, with overall, a greater increase in yield in northern France than in the south. Gross and net primary productivities were more negatively affected by soil water and atmospheric water vapor saturation deficits in western France because of a more pronounced shift in seasonal rainfall from summer to winter. The rotation-averaged values of carbon flux and production for different forest management options were estimated during four years (1980, 2015, 2045 and 2080). Predictions were made using a two-dimensional matrix covering the range of local soil and climate conditions. The changes in ecosystem fluxes and forest production were explained by the counterbalancing effect of rising CO2 concentration and increasing water deficit. The effect of climate change decreased with rotation length from short rotations with high production rates and low standing biomasses to long rotations with low productivities and greater standing biomasses. Climate effects on productivity, both negative and positive, were greatest on high fertility sites. Forest productivity in northern France was enhanced by climate change, increasingly from west to east, whereas in the southwestern Atlantic region, productivity was reduced by climate change to an increasing degree from west to east.

Variability of stem and branch maintenance respiration in a Pinus pinaster tree.

The relationship between maintenance respiration (Rm) of woody organs and their structural characteristics was explored in adult Pinus pinaster Ait. trees. We measured Rm on 75 stem and branch segments of different ages (from 3 to 24 years) and diameters (from 1 to 35 cm). The temperature response of Rm was derived from field measurements based on a classical exponential function with Q10 = 2.13. Relationships between Rm and the dimensions of the woody organs were analyzed under controlled conditions in the laboratory. The surface area of a woody organ was a better predictor of Rm than volume, but surface area failed to account for the observed within-tree variability of Rm among stems, branches and twigs. Two simple models were proposed to predict the variability of Rm at 15 degrees C in an adult tree. Model 1, a linear function model based on the dry mass and nitrogen concentration of sapwood and phloem tissues, explained most of the variability of Rm in branches and stems (R2 = 0.97). We concluded that the respective contributions of the phloem and sapwood depend on the location and diameter of the woody organ. Model 2, a power-law function model based on the length, diameter and age of the sample, explained the same variance of Rm as Model 1 and is appropriate for scaling Rm to the stand level. Models 1 and 2 appear to explain a larger variability of Rm than models based on stem area or sapwood mass.

Combined effects of defoliation and water stress on pine growth and non-structural carbohydrates.

Climate change is expected to increase both pest insect damage and the occurrence of severe drought. There is therefore a need to better understand the combined effects of biotic and abiotic damage on tree growth in order to predict the multi-factorial effect of climate change on forest ecosystem productivity. Indeed, the effect of stress interactions on tree growth is an increasingly important topic that greatly lacks experiments and data, and it is unlikely that the impact of combined stresses can be extrapolated from the outcomes of studies that focused on a single stress. We developed an original manipulative study under real field conditions where we applied artificial defoliation and induced water stress on 10-year-old (∼10 m high) maritime pine trees (Pinus pinaster Ait.). Tree response to combined stresses was quantitatively assessed following tree secondary growth and carbohydrate pools. Such a design allowed us to address the crucial question of combined stresses on trees under stand conditions, sharing soil supplies with neighboring trees. Our initial hypotheses were that (i) moderate defoliation can limit the impact of water stress on tree growth through reduced transpiration demand by a tree canopy partly defoliated and that (ii) defoliation results in reduced non-structural carbohydrate (NSC) pools, affecting tree tolerance to drought. Our results showed additive effects of defoliation and water stress on tree growth and contradict our initial hypothesis. Indeed, under stand conditions, we found that partial defoliation does not limit the impact of water stress through reduced transpiration. Our study also highlighted that, even if NSC in all organs were affected by defoliation, tree response to water stress was not triggered. We found that stem NSC were maintained or increased during the entire growing season, supporting literature-based hypotheses such as an active maintenance of the hydraulic system or another limiting resource for tree growth under defoliation. We also observed a significant decrease in root carbohydrates, which suggests a shift in the root carbon balance under defoliation. The decrease in carbohydrate supply under defoliation may not counterbalance the carbon use for mineral and water uptakes or a translocation to other tissues.

Paired comparison of water, energy and carbon exchanges over two young maritime pine stands (Pinus pinaster Ait.): effects of thinning and weeding in the early stage of tree growth.

The effects of management practices on energy, water and carbon exchanges were investigated in a young pine plantation in south-west France. In 2009-10, carbon dioxide (CO(2)), H(2)O and heat fluxes were monitored using the eddy covariance and sap flow techniques in a control plot (C) with a developed gorse layer, and an adjacent plot that was mechanically weeded and thinned (W). Despite large differences in the total leaf area index and canopy structure, the annual net radiation absorbed was only 4% lower in plot W. We showed that higher albedo in this plot was offset by lower emitted long-wave radiation. Annual evapotranspiration (ET) from plot W was 15% lower, due to lower rainfall interception and transpiration by the tree canopy, partly counterbalanced by the larger evaporation from both soil and regrowing weedy vegetation. The drainage belowground from plot W was larger by 113 mm annually. The seasonal variability of ET was driven by the dynamics of the soil and weed layers, which was more severely affected by drought in plot C. Conversely, the temporal changes in pine transpiration and stem diameter growth were synchronous between sites despite higher soil water content in the weeded plot. At the annual scale, both plots were carbon sinks, but thinning and weeding reduced the carbon uptake by 73%: annual carbon uptake was 243 and 65 g C m(-2) on plots C and W, respectively. Summer drought dramatically impacted the net ecosystem exchange: plot C became a carbon source as the gross primary production (GPP) severely decreased. However, plot W remained a carbon sink during drought, as a result of decreases in both GPP and ecosystem respiration (R(E)). In winter, both plots were carbon sources, plots C and W emitting 67.5 and 32.4 g C m(-2), respectively. Overall, this study highlighted the significant contribution of the gorse layer to mass and energy exchange in young pine plantations.