Repeated summer drought and re-watering during the first growing year of oak (Quercus petraea) delay autumn senescence and bud burst in the following spring

Climate change predicts harsher summer droughts for mid-latitudes in Europe. To enhance our understanding of the putative impacts on forest regeneration, we studied the response of oak seedlings (Quercus petraea) to water deficit. Potted seedlings originating from three locally sourced provenances were subjected to two successive drought periods during the first growing season each followed by a plentiful re watering. Here, we describe survival and phenological responses after the second drought treatment, applying general linear mixed modeling. From the 441 drought treated seedlings 189 subsisted with higher chances of survival among smaller plants and among single plants per pot compared to doubles. Remarkably, survival was independent of the provenance, although relatively more plants had died off in two provenances compared to the third one with mean plant height being higher in one provenance and standard deviation of plant height being higher in the other. Timing of leaf senescence was clearly delayed after the severe drought treatment followed by re-watering, with two seedlings per pot showing a lesser retardation compared to single plants. This delay can be interpreted as a compensation time in which plants recover before entering the subsequent developmental process of leaf senescence, although it renders seedlings more vulnerable to early autumn frosts because of the delayed hardening of the shoots. Onset of bud flush in the subsequent spring still showed a significant but small delay in the drought treated group, independent of the number of seedlings per pot, and can be considered as an after effect of the delayed senescence. In both phenological models significant differences among the three provenances were detected independent from the treatment. The only provenance that is believed to be local of origin, displayed the earliest leaf senescence and the latest flushing, suggesting an adaptation to the local maritime climate. This provenance also displayed the highest standard deviation of plant height, which can be interpreted as an adaptation to variable and unpredictable weather conditions, favoring smaller plants in drought-prone summers and higher plants in more normal growing seasons

Osmolality and non-structural carbohydrate composition in the secondary phloem of trees across a latitudinal gradient in Europe

Phloem osmolality and its components are involved in basic cell metabolism, cell growth, and in various physiological processes including the ability of living cells to withstand drought and frost. Osmolality and sugar composition responses to environmental stresses have been extensively studied for leaves, but less for the secondary phloem of plant stems and branches. Leaf osmotic concentration and the share of pinitol and raffinose among soluble sugars increase with increasing drought or cold stress, and osmotic concentration is adjusted with osmoregulation. We hypothesize that similar responses occur in the secondary phloem of branches. We collected living bark samples from branches of adult Pinus sylvestris, Picea abies, Betula pendula and Populus tremula trees across Europe, from boreal Northern Finland to Mediterranean Portugal. In all studied species, the observed variation in phloem osmolality was mainly driven by variation in phloem water content, while tissue solute content was rather constant across regions. Osmoregulation, in which osmolality is controlled by variable tissue solute content, was stronger for Betula and Populus in comparison to the evergreen conifers. Osmolality was lowest in mid-latitude region, and from there increased by 37% toward northern Europe and 38% toward southern Europe due to low phloem water content in these regions. The ratio of raffinose to all soluble sugars was negligible at mid-latitudes and increased toward north and south, reflecting its role in cold and drought tolerance. For pinitol, another sugar known for contributing to stress tolerance, no such latitudinal pattern was observed. The proportion of sucrose was remarkably low and that of hexoses (i.e., glucose and fructose) high at mid-latitudes. The ratio of starch to all non-structural carbohydrates increased toward the northern latitudes in agreement with the build-up of osmotically inactive C reservoir that can be converted into soluble sugars during winter acclimation in these cold regions. Present results for the secondary phloem of trees suggest that adjustment with tissue water content plays an important role in osmolality dynamics. Furthermore, trees acclimated to dry and cold climate showed high phloem osmolality and raffinose proportion.Peer reviewe

11C-PET imaging reveals transport dynamics and sectorial plasticity of oak phloem after girdling

Carbon transport processes in plants can be followed non-invasively by repeated application of the short-lived positron-emitting radioisotope 11C, a technique which has rarely been used with trees. Recently, positron emission tomography (PET) allowing 3D visualization has been adapted for use with plants. To investigate the effects of stem girdling on the flow of assimilates, leaves on first order branches of two-year-old oak (Quercus robur L.) trees were labeled with 11C by supplying 11CO2-gas to a leaf cuvette. Magnetic resonance imaging gave an indication of the plant structure, while PET registered the tracer flow in a stem region downstream from the labeled branches. After repeated pulse labeling, phloem translocation was shown to be sectorial in the stem: leaf orthostichy determined the position of the phloem sieve tubes containing labeled 11C. The observed pathway remained unchanged for days. Tracer time-series derived from each pulse and analysed with a mechanistic model showed for two adjacent heights in the stem a similar velocity but different loss of recent assimilates. With either complete or partial girdling of bark within the monitored region, transport immediately stopped and then resumed in a new location in the stem cross-section, demonstrating the plasticity of sectoriality. One day after partial girdling, the loss of tracer along the interrupted transport pathway increased, while the velocity was enhanced in a non-girdled sector for several days. These findings suggest that lateral sugar transport was enhanced after wounding by a change in the lateral sugar transport path and the axial transport resumed with the development of new conductive tissue

Early summer drought stress during the first growing year stimulates extra shoot growth in oak seedlings (Quercus petraea)

More severe summer droughts are predicted for mid-latitudes in Europe. To evaluate the impact on forest ecosystems and more specifically on forest regeneration, we studied the response to summer drought in oak seedlings (Quercus petraea). Acorns were collected from different mother trees in three stands in Belgium, sown in pots and grown in non-heated greenhouse conditions. We imposed severe drought on the seedlings in early summer by first watering the pots to saturation and then stopping any watering. Weight of the pots and stomatal conductance were regularly measured. Re-watering followed this drought period of five weeks. Height of the seedlings and apical bud development were observed. Stomatal resistance increased towards the end of the experiment in the drought-treated group and was restored after re-watering. The seedlings from the drought treatment displayed a higher probability to produce additional shoot growth after re-watering (p ≤ 0.05). A higher competition for resources (two plants per pot) increased this chance. Although this chance was also higher for smaller seedlings, the actual length of the extra growth after re-watering was higher for larger seedlings (p ≤ 0.01). Both in the drought-treated and in the control group the autochthonous provenance growing on a xeric site produced less extra shoots compared to the two other provenances. Finally, stressed plants showed less developed apical buds compared to the control group after re-watering, suggesting a phenological effect on the growth cycle of oaks (p ≤ 0.0001). The higher chance for an extra shoot growth after the drought period can be considered as a compensation for the induced growth arrest during the drought period