Twenty years of irrigation acclimation is driven by denser canopies and not by plasticity in twig- and needle-level hydraulics in a Pinus sylvestris forest.

Climate change is predicted to increase atmospheric vapor pressure deficit, exacerbating soil drought, and thus enhancing tree evaporative demand and mortality. Yet, few studies have addressed the longer-term drought acclimation strategy of trees, particularly the importance of morphological versus hydraulic plasticity. Using a long-term (20 years) irrigation experiment in a natural forest, we investigated the acclimation of Scots pine (Pinus sylvestris) morpho-anatomical traits (stomatal anatomy and crown density) and hydraulic traits (leaf water potential, vulnerability to cavitation (Ψ50), specific hydraulic conductivity (Ks), and tree water deficit) to prolonged changes in soil moisture. We found that low water availability reduced twig water potential and increased tree water deficit during the growing season. Still, the trees showed limited adjustments in most branch-level hydraulic traits (Ψ50 and Ks) and needle anatomy. In contrast, trees acclimated to prolonged irrigation by increasing their crown density and hence the canopy water demand. This study demonstrates that despite substantial canopy adjustments, P. sylvestris may be vulnerable to extreme droughts because of limited adjustment potential in its hydraulic system. While sparser canopies reduce water demand, such shifts take decades to occur under chronic water deficits and might not mitigate short-term extreme drought events.

Responses of stem growth and canopy greenness of temperate conifers to dry spells.

Dry spells strongly influence biomass production in forest ecosystems. Their effects may last several years following a drought event, prolonging growth reduction and therefore restricting carbon sequestration. Yet, our understanding of the impact of dry spells on the vitality of trees' above-ground biomass components (e.g., stems and leaves) at a landscape level remains limited. We analyzed the responses of Pinus sylvestris and Picea abies to the four most severe drought years in topographically complex sites. To represent stem growth and canopy greenness, we used chronologies of tree-ring width and time series of the Normalized Difference Vegetation Index (NDVI). We analyzed the responses of radial tree growth and NDVI to dry spells using superposed epoch analysis and further explored this relationship using mixed-effect models. Our results show a stronger and more persistent response of radial growth to dry spells and faster recovery of canopy greenness. Canopy greenness started to recover the year after the dry spell, whereas radial tree growth remained reduced for the two subsequent years and did not recover the pre-drought level until the fourth year after the event. Stem growth and canopy greenness were influenced by climatic conditions during and after drought events, while the effect of topography was marginal. The opposite responses of stem growth and canopy greenness following drought events suggest a different impact of dry spells on trees´ sink and source compartments. These results underscore the crucial importance of understanding the complexities of tree growth as a major sink of atmospheric carbon.

Can mixing Quercus robur and Quercus petraea with Pinus sylvestris compensate for productivity losses due to climate change?

The climate change scenarios RCP 4.5 and RCP 8.5, with a representative concentration pathway for stabilization of radiative forcing of 4.5 W m-2 and 8.5 W m-2 by 2100, respectively, predict an increase in temperature of 1-4.5° Celsius for Europe and a simultaneous shift in precipitation patterns leading to increased drought frequency and severity. The negative consequences of such changes on tree growth on dry sites or at the dry end of a tree species distribution are well-known, but rarely quantified across large gradients. In this study, the growth of Quercus robur and Quercus petraea (Q. spp.) and Pinus sylvestris in pure and mixed stands was predicted for a historical scenario and the two climate change scenarios RCP 4.5 and RCP 8.5 using the individual tree growth model PrognAus. Predictions were made along an ecological gradient ranging from current mean annual temperatures of 5.5-11.4 °C and with mean annual precipitation sums of 586-929 mm. Initial data for the simulation consisted of 23 triplets established in pure and mixed stands of Q. spp. and P. sylvestris. After doing the simulations until 2100, we fitted a linear mixed model using the predicted volume in the year 2100 as response variable to describe the general trends in the simulation results. Productivity decreased for both Q. spp. and P. sylvestris with increasing temperature, and more so, for the warmer sites of the gradient. P. sylvestris is the more productive tree species in the current climate scenario, but the competitive advantage shifts to Q. spp., which is capable to endure very high negative water potentials, for the more severe climate change scenario. The Q. spp.-P. sylvestris mixture presents an intermediate resilience to increased scenario severity. Enrichment of P. sylvestris stands by creating mixtures with Q. spp., but not the opposite, might be a right silvicultural adaptive strategy, especially at lower latitudes. Tree species mixing can only partly compensate productivity losses due to climate change. This may, however, be possible in combination with other silvicultural adaptation strategies, such as thinning and uneven-aged management.

[Temporal Variations in Community Structure and Function of Root-associated Fungi Associated with Pinus sylvestris var. mongholica Plantations in the Mu Us Sandy Land].

To illuminate the temporal variations in the structure and functional groups of the root-associated fungal community associated with Mongolian pine Pinus sylvestris var. mongholica plantations in the Mu Us Sandy Land, P. sylvestris var. mongholica plantations with different stand ages （23, 33, and 44 a） were targeted. The community compositions and main drivers of root-associated fungi at different months and stand ages were identified using the Illumina high-throughput sequencing method. The results indicated that： ① There was a distinct temporal distribution in the root-associated fungal community, the sampling month had a significant effect on the diversity of root-associated fungi （P<0.05）, and the values were higher in May and July. The stand age had no significant effect on the diversity of root-associated fungi （P>0.05） and decreased gradually with increasing stand age. ② The dominant phylum of the root-associated fungal community was Ascomycota. The relative abundance of fungal function groups was different within each month and stand age, and the dominant groups were saprotroph-symbiotroph, undefined saprotroph, and ectomycorrhizal fungi. The indicator genera of ectomycorrhizal fungi in May, July, and September were Melanoleuca, Amphinema, and Tricholoma, respectively. ③ The temporal distribution of the root-associated fungal community was significantly affected by annual relative humidity, annual precipitation, soil porosity, ammonia nitrogen, annual sunshine duration, annual temperature, and soil water content （P<0.05）. Soil organic carbon content, soil porosity, annual precipitation, and annual relative humidity were the main factors that significantly affected the indicator genus of the root-associated fungal community. Our results demonstrated that the temporal distribution of the root-associated fungal community was shaped by climate and soil properties, whereas stand age contributed less. This improved information will provide a theoretical basis for the sustainable management of P. sylvestris var mongholica plantations.

Diverging growth trends and climate sensitivities of individual pine trees after the 1976 extreme drought.

Summer droughts are affecting the productivity and functioning of central European forests, with potentially lasting consequences for species composition and carbon sequestration. Long-term recovery rates and individual growth responses that may diverge from species-specific and population-wide behaviour are, however, poorly understood. Here, we present 2052 pine (Pinus sylvestris) ring width series from 19 forest sites in south-west Germany to investigate growth responses of individual trees to the exceptionally hot and dry summer of 1976. This outstanding drought event presents a distinctive test case to examine long-term post-drought recovery dynamics. We have proposed a new classification approach to identify a distinct sub-population of trees, referred to as "temporarily affected trees", with a prevalence ranging from 9 to 33 % across the forest stands. These trees exhibited an exceptionally prolonged growth suppression, lasting over a decade, indicating significantly lower resilience to the 1976 drought and a 50 % reduced capacity to recover to pre-drought states. Furthermore, shifts in resilience and recovery dynamics are accompanied by changing climate sensitivities, notably an increased response to maximum temperatures and summer droughts in post-1976 affected pines. Our findings underscore the likely interplay between individual factors and micro-site conditions that contribute to divergent tree responses to droughts. Assessing these factors at the individual tree level is recommended to advancing our understanding of forest responses to extreme drought events. By analyzing sub-population growth patterns, our study provides valuable insights into the impacts of summer droughts on central European forests in context of increasing drought events.

Estimation of phloem conductance at tree level in young, middle-aged and old-aged Scots pine trees growing in different climatic conditions in boreal forests.

In forests, a significant proportion of the carbon fixed by trees during photosynthesis is transported belowground along the conducting phloem, so variations in phloem anatomy can lead to variations in transport capacity. Phloem conductance at tree level (Ktree) is also affected by tree height. Both the phloem anatomy and the tree size change during ontogeny, and also differ under different environmental conditions. The goal of our work was to identify the main drivers of variation in Ktree in Scots pine trees growing in natural boreal forests. We conducted a phloem anatomical study and calculated Ktree in trees of three age groups growing in different climatic conditions along a latitudinal gradient from south to north. We found that Ktree was maintained at the same level in actively growing pine trees (25-80-years-old) but increased in old-aged trees (180-190-years-old), possibly reflecting the shift in source-sink relationships of aboveground and belowground parts of trees. Trees of the same age group growing in different climatic conditions demonstrated similar values of Ktree due to coordinated changes in the phloem anatomy and the tree height. In general, the negative influence of tree height on Ktree is offset by the positive influence of phloem width (or trunk diameter) and sieve cell diameter. The exception was young trees growing in the transition zone of the northern taiga subzone to the tundra, where Ktree was the highest in its age group and even exceeded Ktree of middle-aged trees.

Soil C:N:P stoichiometry in rhizosphere and non-rhizosphere of Pinus sylvestris var. mongolica forests.

The aim of this study was to reveal the stoichiometric characteristics of carbon, nitrogen and phosphorus in rhizosphere and non-rhizosphere soils of Pinus sylvestris var. mongolica in the Hulunbuir desert. We investigated the contents and stoichiometry of organic carbon, total nitrogen, and total phosphorus contents of rhizosphere and non-rhizosphere soils across different stand ages (28, 37 and 46 a) of P. sylvestris var. mongolica plantations, with P. sylvestris var. mongolica natural forest as the control. We analyzed the correlation between soils properties and soil stoichiometry. The results showed that rhizosphere effect significantly affected soil N:P, and stand age significantly affected soil organic carbon content in P. sylvestris var. mongolica plantation. Soil organic carbon content in plantation was significantly lower than that in natural forest. Soil organic carbon and total nitrogen contents of plantations in both rhizosphere and non-rhizosphere soils firstly decreased and then increased with increasing stand age, while total phosphorus firstly increased and then decreased in rhizosphere soils, and firstly decreased and then increased in non-rhizosphere soils. There was significant positive correlations between C:N and C:P in rhizosphere soils but not in non-rhizosphere soils, suggesting that higher synergistic rhizosphere soil N and P limitation. The mean N:P values of rhizosphere and non-rhizosphere soils were 4.98 and 8.40, respectively, indicating that the growth of P. sylvestris var. mongolica was restricted by soil N and the rhizosphere soils were more N-restricted. The C:N:P stoichiometry of rhizosphere and non-rhizosphere soils were significantly influenced by soil properties, with available phosphorus being the most important driver. The growth of P. sylvestris var. mongolica was limited by N in the Hulunbuir desert, and root system played an obvious role in enriching and maintaining soil nutrients. It was recommended that soil nitrogen should be supplemented appropriately during the growth stage of P. sylvestris var. mongolica plantation, and phosphorus should be supplemented appropriately according to the synergistic nature of nitrogen and phosphorus limitation.