Research frontiers for improving our understanding of drought-induced tree and forest mortality.

Accumulating evidence highlights increased mortality risks for trees during severe drought, particularly under warmer temperatures and increasing vapour pressure deficit (VPD). Resulting forest die-off events have severe consequences for ecosystem services, biophysical and biogeochemical land-atmosphere processes. Despite advances in monitoring, modelling and experimental studies of the causes and consequences of tree death from individual tree to ecosystem and global scale, a general mechanistic understanding and realistic predictions of drought mortality under future climate conditions are still lacking. We update a global tree mortality map and present a roadmap to a more holistic understanding of forest mortality across scales. We highlight priority research frontiers that promote: (1) new avenues for research on key tree ecophysiological responses to drought; (2) scaling from the tree/plot level to the ecosystem and region; (3) improvements of mortality risk predictions based on both empirical and mechanistic insights; and (4) a global monitoring network of forest mortality. In light of recent and anticipated large forest die-off events such a research agenda is timely and needed to achieve scientific understanding for realistic predictions of drought-induced tree mortality. The implementation of a sustainable network will require support by stakeholders and political authorities at the international level.

Fate of xylem-transported 11C- and 13C-labeled CO2 in leaves of poplar.

In recent studies, assimilation of xylem-transported CO2 has gained considerable attention as a means of recycling respired CO2 in trees. However, we still lack a clear and detailed picture on the magnitude of xylem-transported CO2 assimilation, in particular within leaf tissues. To this end, detached poplar leaves (Populus × canadensis Moench 'Robusta') were allowed to take up a dissolved (13)CO2 label serving as a proxy of xylem-transported CO2 entering the leaf from the branch. The uptake rate of the (13)C was manipulated by altering the vapor pressure deficit (VPD) (0.84, 1.29 and 1.83 kPa). Highest tissue enrichments were observed under the highest VPD. Among tissues, highest enrichment was observed in the petiole and the veins, regardless of the VPD treatment. Analysis of non-labeled leaves showed that some (13)C diffused from the labeled leaves and was fixed in the mesophyll of the non-labeled leaves. However, (13)C leaf tissue enrichment analysis with elemental analysis coupled to isotope ratio mass spectrometry was limited in spatial resolution at the leaf tissue level. Therefore, (11)C-based CO2 labeling combined with positron autoradiography was used and showed a more detailed spatial distribution within a single tissue, in particular in secondary veins. Therefore, in addition to (13)C, (11) C-based autoradiography can be used to study the fate of xylem-transported CO2 at leaf level, allowing the acquisition of data at a yet unprecedented resolution.

Stem girdling affects the quantity of CO2 transported in xylem as well as CO2 efflux from soil.

There is recent clear evidence that an important fraction of root-respired CO2 is transported upward in the transpiration stream in tree stems rather than fluxing to the soil. In this study, we aimed to quantify the contribution of root-respired CO2 to both soil CO2 efflux and xylem CO2 transport by manipulating the autotrophic component of belowground respiration. We compared soil CO2 efflux and the flux of root-respired CO2 transported in the transpiration stream in girdled and nongirdled 9-yr-old oak trees (Quercus robur) to assess the impact of a change in the autotrophic component of belowground respiration on both CO2 fluxes. Stem girdling decreased xylem CO2 concentration, indicating that belowground respiration contributes to the aboveground transport of internal CO2 . Girdling also decreased soil CO2 efflux. These results confirmed that root respiration contributes to xylem CO2 transport and that failure to account for this flux results in inaccurate estimates of belowground respiration when efflux-based methods are used. This research adds to the growing body of evidence that efflux-based measurements of belowground respiration underestimate autotrophic contributions.

Transport of root-respired CO2 via the transpiration stream affects aboveground carbon assimilation and CO2 efflux in trees.

Upward transport of CO2 via the transpiration stream from belowground to aboveground tissues occurs in tree stems. Despite potentially important implications for our understanding of plant physiology, the fate of internally transported CO2 derived from autotrophic respiratory processes remains unclear. We infused a ¹³CO2-labeled aqueous solution into the base of 7-yr-old field-grown eastern cottonwood (Populus deltoides) trees to investigate the effect of xylem-transported CO2 derived from the root system on aboveground carbon assimilation and CO2 efflux. The ¹³C label was transported internally and detected throughout the tree. Up to 17% of the infused label was assimilated, while the remainder diffused to the atmosphere via stem and branch efflux. The largest amount of assimilated ¹³C was found in branch woody tissues, while only a small quantity was assimilated in the foliage. Petioles were more highly enriched in ¹³C than other leaf tissues. Our results confirm a recycling pathway for respired CO2 and indicate that internal transport of CO2 from the root system may confound the interpretation of efflux-based estimates of woody tissue respiration and patterns of carbohydrate allocation.

Assimilation of xylem-transported CO2 is dependent on transpiration rate but is small relative to atmospheric fixation.

The effect of transpiration rate on internal assimilation of CO2 released from respiring cells has not previously been quantified. In this study, detached branches of Populus deltoides were allowed to take up (13)CO2-labelled solution at either high (high label, HL) or low (low label, LL) (13)CO2 concentrations. The uptake of the (13)CO2 label served as a proxy for the internal transport of respired CO2, whilst the transpiration rate was manipulated at the leaf level by altering the vapour pressure deficit (VPD) of the air. Simultaneously, leaf gas exchange was measured, allowing comparison of internal CO2 assimilation with that assimilated from the atmosphere. Subsequent (13)C analysis of branch and leaf tissues revealed that woody tissues assimilated more label under high VPD, corresponding to higher transpiration, than under low VPD. More (13)C was assimilated in leaf tissue than in woody tissue under the HL treatment, whereas more (13)C was assimilated in woody tissue than in leaf tissue under the LL treatment. The ratio of (13)CO2 assimilated from the internal source to CO2 assimilated from the atmosphere was highest for the branches under the HL and high VPD treatment, but was relatively small regardless of VPD×label treatment combination (up to 1.9%). These results showed that assimilation of internal CO2 is highly dependent on the rate of transpiration and xylem sap [CO2]. Therefore, it can be expected that the relative contribution of internal CO2 recycling to tree carbon gain is strongly dependent on factors controlling transpiration, respiration, and photosynthesis.

Water use of a multigenotype poplar short-rotation coppice from tree to stand scale.

Short-rotation coppice (SRC) has great potential for supplying biomass-based heat and energy, but little is known about SRC's ecological footprint, particularly its impact on the water cycle. To this end, we quantified the water use of a commercial scale poplar (Populus) SRC plantation in East Flanders (Belgium) at tree and stand level, focusing primarily on the transpiration component. First, we used the AquaCrop model and eddy covariance flux data to analyse the different components of the stand-level water balance for one entire growing season. Transpiration represented 59% of evapotranspiration (ET) at stand scale over the whole year. Measured ET and modelled ET were lower as compared to the ET of reference grassland, suggesting that the SRC only used a limited amount of water. Secondly, we compared leaf area scaled and sapwood area scaled sap flow (Fs) measurements on individual plants vs. stand scale eddy covariance flux data during a 39-day intensive field campaign in late summer 2011. Daily stem diameter variation (∆D) was monitored simultaneously with Fs to understand water use strategies for three poplar genotypes. Canopy transpiration based on sapwood area or leaf area scaling was 43.5 and 50.3 mm, respectively, and accounted for 74%, respectively, 86%, of total ecosystem ET measured during the intensive field campaign. Besides differences in growth, the significant intergenotypic differences in daily ∆D (due to stem shrinkage and swelling) suggested different water use strategies among the three genotypes which were confirmed by the sap flow measurements. Future studies on the prediction of SRC water use, or efforts to enhance the biomass yield of SRC genotypes, should consider intergenotypic differences in transpiration water losses at tree level as well as the SRC water balance at stand level.

Internal recycling of respired CO 2 may be important for plant functioning under changing climate regimes.

Recent studies have provided evidence of a large flux of root-respired CO 2 in the transpiration stream of trees. In our study, we investigated the potential impact of this internal CO 2 transport on aboveground carbon assimilation and CO 2 efflux. To trace the transport of root-respired CO 2, we infused a (13)C label at the stem base of field-grown Populus deltoides Bartr. ex. Marsh trees. The (13)C label was transported to the top of the stem and throughout the crown via the transpiration stream. Up to 17% of the (13)C label was assimilated by chlorophyll-containing tissues. Our results provide evidence of a mechanism for recycling respired CO 2 within trees. Such a mechanism may have important implications for how plants cope with predicted increases in intensity and frequency of droughts. Here, we speculate on the potential significance of this recycling mechanism within the context of plant responses to climate change and plants currently inhabiting arid environments.