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Aims Fine roots, that comprise the adjustable part of the root system, are important in spatially heterogeneous boreal forest soils. We investigated the soil exploring patterns of Norway spruce (Picea abies) seedlings of equal height belonging to families representing two contrasting growth phenotypes that have shown fast and slow growth rates in long-term experiments. We hypothesised that seedlings of the fast-growing phenotype would show a more explorative root growth strategy, intense branching, and root proliferation in response to the nutrient patch, and that slow-growing phenotypes would be more tolerant to drought stress. Methods Seedlings were grown in flat Perspex microcosms with a clod of humus placed in the mid-bottom part of each microcosm for eight months. The order-based and functional classification, branching topology, and size of seedling root systems were studied with WinRHIZO™ image-analysis software and root exploration patterns with LIGNUM-model simulations. In addition, transpiration, stomatal conductance, net assimilation rate responses were measured. Results No differences were found in the early foraging of roots for the humus clod nor net assimilation rate and transpiration between the phenotype groups. Seedlings were favouring exploitation over exploration in the early phases of development regardless of growth phenotype group. However, in fast-growing phenotypes, the main roots were longer, and the lateral root pool favoured long and bifurcated laterals that formed larger absorptive root area. Conclusions Our results indicate that in nutrient-poor conditions, better growth of lateral roots precedes future differences in the aboveground growth rate of Norway spruce.

Urban green areas are becoming increasingly recognized for their biodiversity potential. However, little is known about how urbanization shapes cryptic species communities, such as those residing in deadwood. In this study, we investigated downed Norway spruce trunks at intermediate stages of decay, in urban and semi-natural forests in southern Finland. To understand the interconnections between landscape context, deadwood characteristics and wood-inhabiting fungal communities, we studied structural characteristics, surface epiphyte cover and internal moisture and temperature conditions of the tree trunks, and fungal communities residing in the wood. Our findings showed that urban tree trunks had less epiphyte cover and lower moisture than trunks in semi-natural forests. Overall, urban forests provide less favourable habitats for a majority of the dominant wood-inhabiting fungal species and for red-listed species as a group. Yet, 33% of urban trunks hosted at least one red-listed species. While these landscape-scale effects may be driven by local climatic conditions as well as contingencies related to available species pools, our results also highlight the significance of substrate-scale variability of deadwood in shaping wood-inhabiting fungal communities. We show that epiphyte cover is a significant driver or indicator of these small-scale dynamic processes in deadwood.

Trees in northern latitude ecosystems are projected to experience increasing drought stress as a result of rising air temperatures and changes in precipitation patterns in northern latitude ecosystems. However, most drought-related studies on high-latitude boreal forests (>50 degrees N) have been conducted in North America, with few studies quantifying the response in European and Eurasian boreal forests. Here, we tested how daily whole-tree transpiration (Q, Liters day(-1)) and Q normalized for mean daytime vapor pressure deficit (Q(DZ), Liters day(-1) kPa(-1)) were affected by the historic 2018 drought in Europe. More specifically, we examined how tree species, size, and topographic position affected drought response in high-latitude mature boreal forest trees. We monitored 30 Pinus sylvestris (pine) and 30 Picea abies (spruce) trees distributed across a topographic gradient in northern Sweden. In general, pine showed a greater Q(DZ) control compared to spruce during periods of severe drought (standardized precipitation-evapotranspiration index: SPEI

Plant respiratory burst oxidase homologs (RBOHs) are plasma membrane-localized NADPH oxidases that generate superoxide anion radicals, which then dismutate to H2O2, into the apoplast using cytoplasmic NADPH as an electron donor. PaRBOH1 is the most highly expressed RBOH gene in developing xylem as well as in a lignin-forming cell culture of Norway spruce (Picea abies L. Karst.). Since no previous information about regulation of gymnosperm RBOHs exist, our aim was to resolve how PaRBOH1 is regulated with a focus on phosphorylation. The N-terminal part of PaRBOH1 was found to contain several putative phosphorylation sites and a four-times repeated motif with similarities to the Botrytis-induced kinase 1 target site in Arabidopsis AtRBOHD. Phosphorylation was indicated for six of the sites in in vitro kinase assays using 15 amino-acid-long peptides for each of the predicted phosphotarget site in the presence of protein extracts of developing xylem. Serine and threonine residues showing positive response in the peptide assays were individually mutated to alanine (kinase-inactive) or to aspartate (phosphomimic), and the wild type PaRBOH1 and the mutated constructs transfected to human kidney embryogenic (HEK293T) cells with a low endogenous level of extracellular ROS production. ROS-producing assays with HEK cells showed that Ca2+ and phosphorylation synergistically activate the enzyme and identified several serine and threonine residues that are likely to be phosphorylated including a novel phosphorylation site not characterized in other plant species. These were further investigated with a phosphoproteomic study. Results of Norway spruce, the first gymnosperm species studied in relation to RBOH regulation, show that regulation of RBOH activity is conserved among seed plants.

Embryogenic tissues of Abies alba Mill. were cryopreserved using the slow-freezing approach. Four cell lines were incubated for 24 h on a medium with 0.5 M sorbitol and pre-treated with 5% DMSO. Subsequently, the tissues were frozen at a cooling rate of 1 °C min-1 to -40 °C and transferred to liquid nitrogen for 72 hours. After thawing in a water bath at 40 °C, the tissues were cultivated on a proliferation medium. All tested lines recovered, but variations in regrowth frequencies across cell lines were noticed (91.66 to 100%). The recovered tissues showed similar features to the control 2 (non-pre-treated and non-cryopreserved tissues). In the accumulation of fresh and dry mass, no statistically significant differences were observed between cryopreserved cultures and control 2. The cryopreserved tissues produced cotyledonary somatic embryos capable of germination. Microscopic observations revealed considerable structural changes as a consequence of the cryopreservation procedure. The long vacuolated suspensor cells were disrupted, and mostly the meristematic cells of the embryonal region survived. The typical bipolar structure of early somatic embryos has been regained during the post-thaw period. Differences in cryotolerance across cell lines were also observed.

Tree stems are an overlooked source of volatile organic compounds (VOCs). Their contribution to ecosystem processes and total VOC fluxes is not well studied, and assessing it requires better understanding of stem emission dynamics and their driving processes. To gain more mechanistic insight into stem emission patterns, we measured monoterpene, methanol, and acetaldehyde emissions from the stems of mature Scots pines (Pinus sylvestris L.) in a boreal forest over three summers. We analysed the effects of temperature, soil water content, tree water status, transpiration, and growth on the VOC emissions, and used generalized linear models to test their relative importance in explaining the emissions. We show that Scots pine stems are considerable sources of monoterpenes, methanol, and acetaldehyde, and their emissions are strongly regulated by temperature. However, even small changes in water availability affected the emission potentials: increased soil water content increased the monoterpene emissions within a day, whereas acetaldehyde and methanol emissions responded within two to four days. This lag corresponded to their transport time in the xylem sap from the roots to the stem. Moreover, the emissions of monoterpenes, methanol, and acetaldehyde were influenced by the cambial growth rate of the stem with six- to ten-day lags. This article is protected by copyright. All rights reserved.

Extreme droughts are expected to increase in frequency and severity in many regions of the world, threatening multiple ecosystem services provided by forests. Effective strategies to adapt forests to such droughts require comprehensive information on the effects and importance of the factors influencing forest resistance and resilience. We used a unique combination of inventory and dendrochronological data from a long‐term (>30 years) silvicultural experiment in mixed silver fir and Norway spruce mountain forests along a temperature and precipitation gradient in southwestern Germany. We aimed at examining the mechanisms and forest stand characteristics underpinning the resistance and resilience to past mild and severe droughts. We found that (i) fir benefited from mild droughts and showed higher resistance (i.e., lower growth loss during drought) and resilience (i.e., faster return to pre‐drought growth levels) than spruce to all droughts; (ii) species identity determined mild drought responses while species interactions and management‐related factors strongly influenced the responses to severe droughts; (iii) intraspecific and interspecific interactions had contrasting effects on the two species, with spruce being less resistant to severe droughts when exposed to interaction with fir and beech; (iv) higher values of residual stand basal area following thinning were associated with lower resistance and resilience to severe droughts; and (v) larger trees were resilient to mild drought events but highly vulnerable to severe droughts. Our study provides an analytical approach for examining the effects of different factors on individual tree‐ and stand‐level drought response. The forests investigated here were to a certain extent resilient to mild droughts, and even benefited from such conditions, but were strongly affected by severe droughts. Lastly, negative effects of severe droughts can be reduced through modifying species composition, tree size distribution and stand density in mixed silver fir‐Norway spruce forests., We examined mechanisms and stand characteristics underpinning the growth resistance (Rt), recovery (Rc) and resilience (Rs) to past mild and severe droughts in mixed silver fir and Norway spruce mountain forests in southwestern Germany. We found that the forests investigated were to a certain extent resilient to mild droughts, and even benefited from such conditions, but were strongly affected by severe droughts (especially those stands with higher residual basal area and larger trees). Lastly, negative effects of severe droughts can be reduced through modifying species composition, tree size distribution and stand density in mixed silver fir‐Norway spruce forests.

The collection of modern, spatially extensive pollen data is important for the interpretation of fossil pollen assemblages and the reconstruction of past vegetation communities in space and time. Modern datasets are readily available for percentage data but lacking for pollen accumulation rates (PARs). Filling this gap has been the motivation of the pollen monitoring network, whose contributors monitored pollen deposition in modified Tauber traps for several years or decades across Europe. Here we present this monitoring dataset consisting of 351 trap locations with a total of 2742 annual samples covering the period from 1981 to 2017. This dataset shows that total PAR is influenced by forest cover and climate parameters, which determine pollen productivity and correlate with latitude. Treeless vegetation produced PAR values of at least 140 grains cm−2 yr−1. Tree PAR increased by at least 400 grains cm−2 yr−1 with each 10 % increase in forest cover. Pollen traps situated beyond 200 km of the distribution of a given tree species still collect occasional pollen grains of that species. The threshold of this long-distance transport differs for individual species and is generally below 60 grains cm−2 yr−1. Comparisons between modern and fossil PAR from the same regions show similar values. For temperate taxa, modern analogues for fossil PARs are generally found downslope or southward of the fossil sites. While we do not find modern situations comparable to fossil PAR values of some taxa (e.g. Corylus), CO2 fertilization and land use may cause high modern PARs that are not documented in the fossil record. The modern data are now publicly available in the Neotoma Paleoecology Database and aid interpretations of fossil PAR data.

Forest fire risk in Finland is estimated with the Finnish Forest Fire Index (FFI), which predicts the fuel moisture content (FMC) of the forest floor. We studied the FMC variation of four typical ground vegetation fuels, Pleurozium schreberi, Hylocomium splendens, Dicranum spp., and Cladonia spp., and raw humus in mature and recently clear-cut stands. Of these, six were sub-xeric Pinus sylvestris stands, and six mesic Picea abies stands. We analysed the ability of the FFI to predict FMC and compared it with the widely applied Canadian Fire Weather Index (FWI). We found that in addition to stand characteristics, ground layer FMC was highly dependent on the species so that Dicranum was the moistest, and Cladonia the driest. In the humus layer, the differences among species were small. Overall, the FWI was a slightly better predictor of FMC than the FFI. While the FFI generally predicted ground layer FMC well, the shape of the relationship varied among the four species. The use of auxiliary variables thus has potential in improving predictions of ignitions and forest fire risk. Knowledge of FMC variation could also benefit planning and timing of prescribed burns.

The collection of modern, spatially extensive pollen data is important for the interpretation of fossil pollen assemblages and the reconstruction of past vegetation communities in space and time. Modern datasets are readily available for percentage data but lacking for pollen accumulation rates (PARs). Filling this gap has been the motivation of the pollen monitoring network, whose contributors monitored pollen deposition in modified Tauber traps for several years or decades across Europe. Here we present this monitoring dataset consisting of 351 trap locations with a total of 2742 annual samples covering the period from 1981 to 2017. This dataset shows that total PAR is influenced by forest cover and climate parameters, which determine pollen productivity and correlate with latitude. Treeless vegetation produced PAR values of at least 140 grains cm(-2) yr(-1). Tree PAR increased by at least 400 grains cm(-2) yr(-1) with each 10% increase in forest cover. Pollen traps situated beyond 200 km of the distribution of a given tree species still collect occasional pollen grains of that species. The threshold of this long-distance transport differs for individual species and is generally below 60 grains cm(-2) yr(-1). Comparisons between modern and fossil PAR from the same regions show similar values. For temperate taxa, modern analogues for fossil PARs are generally found downslope or southward of the fossil sites. While we do not find modern situations comparable to fossil PAR values of some taxa (e.g. Corylus), CO2 fertilization and land use may cause high modern PARs that are not documented in the fossil record. The modern data are now publicly available in the Neotoma Paleoecology Database and aid interpretations of fossil PAR data. Czech Science Foundation (GACR)Grant Agency of the Czech Republic [17-17909S]; Czech Academy of SciencesCzech Academy of Sciences [RVO 67985939]; Charles University Research Centre [204069]; ETAG [PUT323] This research has been supported by the Czech Science Foundation (GACR) (grant no. 17-17909S), the Czech Academy of Sciences (grant no. RVO 67985939), the Charles University Research Centre (grant no. 204069), and the ETAG (grant no. PUT323).

In cut-to-length logging, the harvester operator adjusts the bucking in accordance with visible defects on processed stems. Some of the defects, such as a sweep on the bottom of the stem, decrease the yield and quality of sawn products and are difficult for the operator to notice. Detecting the defects with improved sensors would support the operator in his qualitative decision-making and increase value recovery of logging. Predicting the maximum bow height of the bottom log in Norway spruce (Picea abies (L.) Karst.) with log end face image and stem taper was investigated with two modelling approaches. A total of 101 stems were selected from five clear-cut stands in southern Finland. The stems were crosscut and taper measured, and the butt ends of the bottom logs were photographed. The stem diameter, out-of-roundness, and pith eccentricity were measured from the images while the max. bow height was measured by a 3D log scanner at a sawmill. The bottom logs with an eccentric pith had higher max. bow height. In addition, a highly conical bottom part of the stem was more common on the bottom logs with a large max. bow height. Applying both log end face image and stem taper measurements gave the best model fit and detection accuracy (76%) for bottom logs with a large max. bow height. The results indicate that the log end face image and stem taper measurements can be utilised to aid harvester operator in deciding an optimised length for logs according to the bow height.

Developing shoots, i.e., sprouts, and older needles of Norway spruce (Picea abies (L.) Karst.) have traditionally been used for medicinal purposes due to the high content of vitamins and antioxidants. Currently, sprouts are available as, for example, superfood and supplements. However, end-product quality and nutritive value may decline in the value-chain from raw material sourcing to processing and storage. We studied (1) impacts of different drying and extraction methods on nutritional composition and antioxidative properties of sprouts and needles, (2) differences between sprouts and needles in nutritional composition and microbiological quality, and (3) production scale quality of the sprouts. Additionally, (4) sprout powder was applied in products (ice-cream and sorbet) and consumer acceptance was evaluated. According to our results, older needles have higher content of dry matter, energy, and calcium, but lower microbial quality than sprouts. Sprouts showed a higher concentration of vitamin C, magnesium, potassium, and phosphorus than older needles. Freeze-drying was the best drying method preserving the quality of both sprouts and needles, e.g., vitamin C content. The antioxidative activity of the sprout extracts were lower than that of needles. Ethanol-water extraction resulted in a higher content of active compounds in the extract than water extraction. Sensory evaluation of food products revealed that on average, 76% of consumers considered sprout-containing products very good or good, and a creamy product was preferred over a water-based sorbet.

Bark of Norway spruce and Scots pine trees contain large amounts of condensed tannins. Tannins extracted with hot water could be used in different applications as they possess antioxidative and antimicrobial activities. The use of bark tannins as e.g., food preservatives calls for increases in our knowledge of their antioxidative activities when applied in foodstuffs. To assess the ability of bark tannins to prevent lipid oxidation, hot water extracts were evaluated in a liposome model. Isolated tannins were also applied in dry-cured, salty meat snacks either as liquid extracts or in dry-powder form. Consumer acceptance of the snacks was tested by a sensory evaluation panel where outlook, odor, taste, and structure of the snacks were evaluated and compared to a commercial product without tannin ingredients. Our results show that conifer bark tannin-rich extracts have high capacity to prevent lipid oxidation in the liposome model. The efficacies of pine and spruce bark extracts were ten to hundred folds higher, respectively, than those of phenolic berry extracts. The bark extracts did not significantly influence the odor or taste of the meat snacks. The findings indicate that bark extracts may be used as sustainable food ingredients. However, more research is needed to verify their safety.

Human-induced fragmentation affects forest continuity, i.e. availability of a suitable habitat for the target species over a time period. The dependence of wood-inhabiting fungi on landscape level continuity has been well demonstrated, but the importance of local continuity has remained controversial. In this study, we explored the effects of local forest continuity (microhabitat and stand level) on the diversity of wood-inhabiting fungi on standing dead trunks of Scots pine (Pinus sylvestris L.). We studied species richness and community composition of decomposers and Micarea lichens on 70 trunks in 14 forests in central Finland that differed in their state of continuity. We used dendrochronological methods to assess the detailed history of each study trunk, i.e. the microhabitat continuity. The stand continuity was estimated as dead wood diversity and past management intensity (number of stumps). We recorded 107 species (91 decomposers, 16 Micarea lichens), with a total of 510 occurrences. Using generalized linear mixed models, we found that none of the variables explained decomposer species richness, but that Micarea species richness was positively dependent on the time since tree death. Dead wood diversity was the most important variable determining the composition of decomposer communities. For Micarea lichens, the community composition was best explained by the combined effect of years from death, site and dead wood diversity. However, these effects were rather tentative. The results are in line with those of previous studies suggesting the restricted significance of local forest continuity for wood-inhabiting fungi. However, standing dead pines that have been available continuously over long periods seem to be important for species-rich communities of Micarea lichens. Rare specialists (e.g. on veteran trees) may be more sensitive to local continuity, and should be at the center of future research. peerReviewed

We present spring and summer volatile organic compound (VOC) emission rate measurements from Norway spruce (Picea abies L. Karst) growing in a boreal forest in southern Finland. The measurements were conducted using in situ gas chromatograph with 1 to 2 h time resolution to reveal quantitative and qualitative short-term and seasonal variability of the emissions. The measurements cover altogether 14 weeks in years 2011, 2014 and 2015. Monoterpene (MT) and sesquiterpene (SQT) emission rates were measured all the time, but isoprene only in 2014 and 2015 and acetone and C4–C10 aldehydes only in 2015. The emission rates of all the compounds were low in spring, but MT, acetone, and C4–C10 aldehyde emission rates increased as summer proceeded, reaching maximum emission rates in July. Late summer mean values (late July and August) were 29, 17, and 33 ng g(dw)−1 h−1 for MTs, acetone, and aldehydes respectively. SQT emission rates increased during the summer and highest emissions were measured in late summer (late summer mean value 84 ng g(dw)−1 h−1) concomitant with highest linalool emissions most likely due to stress effects. The between-tree variability of emission pattern was studied by measuring seven different trees during the same afternoon using adsorbent tubes. Especially the contributions of limonene, terpinolene, and camphene were found to vary between trees, whereas proportions of α-pinene (25 ± 5 %) and β-pinene (7 ± 3 %) were more stable. Our results show that it is important to measure emissions at canopy level due to irregular emission pattern, but reliable SQT emission data can be measured only from enclosures. SQT emissions contributed more than 90 % of the ozone reactivity most of the time, and about 70 % of the OH reactivity during late summer. The contribution of aldehydes to OH reactivity was comparable to that of MT during late summer, 10–30 % most of the time.

Beavers are ecosystem engineers that modify and maintain a range of special habitat types in boreal forests. They also produce large quantities of deadwood that provide substrate for many lignicolous organisms such as calicioid fungi (Ascomycota). We studied how calicioid diversity differed between boreal riparian forests with and without beaver activity. The results show that calicioid diversity were significantly higher at beaver sites compared to the other two forest site types studied. The large quantity and diverse forms of deadwood produced by beavers clearly promotes calicioid diversity in the boreal landscape. The specific lighting and humidity conditions within beaver wetlands could be the reason why they promote the success of certain calicioid species.

The artificial recharge of groundwater by infiltrating surface water through forest soil has been introduced as a groundwater producing practice in Finland. As a result, the forest soil, as well as the whole ecosystem, is subjected to extremely high inputs of carbon and nutrient rich lake water. The effects of sprinkling infiltration on forest soil, tree growth and understory vegetation and their respective recovery were studied on a forested esker in central Finland. The Scots pine-dominated experimental plots were sprinkled with lake water in 1998-2001 and sampled after a 12-15-year recovery period. Soil pH and base cation concentration, as well as the rate of net N mineralization were significantly higher at the plots that had been infiltrated. The concentrations of base cations calcium and magnesium were thousands of times higher in the infiltrated soil than in the untreated soil. In addition, sprinkling infiltration had favored early-successional herbs, grasses and forbs and negatively affected late successional, slow-growing mosses and lichens. Sprinkling infiltration had significantly increased tree radial growth. Sprinkling infiltration is an environment altering soil treatment method which, based on the findings of this study, can have long-term effects on tree growth, soil processes and understory vegetation.

Fire is a major disturbance agent in the boreal forest, influencing many current and future ecosystem conditions and services. Surprisingly few studies have attempted to improve the accuracy of fire-event reconstructions even though the estimates of the occurrence of past fires may be biased, influencing the reliability of the models employing those data (e.g. C stock, cycle). This study aimed to demonstrate how three types of fire proxies – fire scars from tree rings, sedimentary charcoal and, for the first time in this context, fungal spores of Neurospora – can be integrated to achieve a better understanding of past fire dynamics. By studying charcoal and Neurospora from sediment cores from forest hollows, and the fire scars from tree rings in their surroundings in the southern Fennoscandian and western Russian boreal forest, we produced composite fire-event data sets and fire-event frequencies, and estimated fire return intervals. Our estimates show that the fire return interval varied between 126 and 237 years during the last 11,000 years. The highest fire frequency during the 18th–19th century can be associated with the anthropogenic influence. Importantly, statistical tests revealed a positive relationship between other fire event indicators and Neurospora occurrence allowing us to pinpoint past fire events at times when the sedimentary charcoal was absent, but Neurospora were abundant. We demonstrated how fire proxies with different temporal resolution can be linked, providing potential improvements in the reliability of fire history reconstructions from multiple proxies.

Information regarding the nature and rate of forest recovery is required to inform forest management, monitoring, and reporting activities. Delayed establishment or return of forests has implications to harvest rotations and carbon uptake, among others, creating a need for spatially-explicit, large-area, characterizations of forest recovery. Landsat time series (LTS) has been demonstrated as a means to quantitatively relate forest recovery, noting that there are gaps in our understanding of the linkage between spectral measures of forest recovery and manifestations of forest structure and composition. Field plots provide a means to better understand the linkage between forest characteristics and spectral recovery indices. As such, from a large set of existing field plots, we considered the conditions present for the year in which the co-located pixel was considered spectrally recovered using the Years to Recovery (Y2R) metric. Y2R is a long-term metric of spectral recovery that indicates the number of years required for a pixel to return to 80% of its pre-disturbance Normalized Burn Ratio value. Absolute and relative metrics of recovery at 5 years post-disturbance were also considered. We used these three spectral recovery metrics to predict the stand development class assigned by the field crew for 284 seedling plots with an overall accuracy of 73.59%, with advanced seedling stands more accurately discriminated (omission error, OE = 15.74%) than young seedling stands (OE = 49.84%). We then used field-measured attributes (e.g. height, stem density, dominant species) from the seedling plots to classify the plots into three spectral recovery groups, which were defined using the Y2R metric: spectral recovery in (1) 1–5 years, (2) 6–10 years, or (3) 11–15 years. Overall accuracy for spectral recovery groups was 61.06%. Recovery groups 1 and 3 were discriminated with greater accuracy (producer’s and user’s accuracies > 66%) than recovery group 2 ( 66%) than recovery group 2 ( 66%) than recovery group 2 (

Phenolics and extracted phenolic compounds of Scots pine (Pinus sylvestris) and Norway spruce (Picea abies) show antibacterial activity against several bacteria. The majority of phenolic compounds are stilbenes, flavonoids, proanthocyanidins, phenolic acids, and lignans that are biosynthesized in the wood through the phenylpropanoid pathway. In Scots pine (P. sylvestris), the most abundant phenolic and antibacterial compounds are pinosylvin-type stilbenes and flavonol- and dihydroflavonol-type flavonoids, such as kaempferol, quercetin, and taxifolin and their derivatives. In Norway spruce (P. abies) on the other hand, the main stilbene is resveratrol and the major flavonoids are quercetin and myricetin. In general, when the results from the literature regarding the activities of flavonoid glycosides and their aglycones against a total of twenty-one microorganisms are summarized, it was found that phenolic glycosides are less active than the corresponding aglycones, although a number of exceptions are also known. The aglycones in plants respond to various kinds of biotic stress. Synergistic effects between aglycones and their glycosides have been observed. Minimum inhibition concentrations of below 10 mg L−1 against bacteria have been reported for gallic acid, apigenin, and several methylated and acylated flavonols present in these industrially important trees. In general, the phenolic compounds are more active against Gram-positive bacteria, but apigenin is reported to exhibit strong activity against Gram-negative bacteria. The present review lists some of the biosynthesis pathways for the antibacterial phenolic metabolites found in Scots pine (P. sylvestris) and Norway spruce (P. abies). The antimicrobial activity of the compounds is collected and compared to gather information about the most effective secondary metabolites.

As climate change should lead to an increase in the vulnerability and the sensitivity of forests to extreme climatic events, quantifying and predicting their response to more severe droughts remains a key task for foresters. Furthermore, recent works have suggested that tree diversity may affect forest ecosystem functioning, including their response to extreme events. In this study we aimed at testing whether the growth response of forest stands to stressful climatic events varied between mixed and monospecific stands, under various environmental conditions. We focused on beech-fir forests (Fagus sylvatica [L.] and Abies alba [L.]) and beech-oak forests (F. sylvatica [L.] and Quercus pubescent [L.]) in the French Alps. We used a dendrochronological dataset sampled in forest plots organized by triplets (one mixture and two monospecific stands) distributed in six sites along a latitudinal gradient. We tested (1) whether stand diversity (two-species stands vs monospecific stands) modulates the stands’ response to drought events in terms of productivity, (2) whether species identity may drive the diversity effect on resistance and recovery, and (3) whether this can be explained by interspecific interactions. We found that (1) interspecific differences in response to extreme drought events (possibly due to interspecific differences in hydraulic characteristics) can induce a mixture effect on stand growth, although it appeared (2) to be strongly depending on species identity (positive effect only found for beech-fir mixed stands), while (3) there were no significant non-additive effects of diversity on stand resistance and recovery, except for some specific cases. Overall, our study shows that promoting selected mixed stands management may buffer extreme drought effect on stand productivity.

Phloem failure has recently been recognized as one of the mechanisms causing tree mortality under drought, though direct evidence is still lacking. We combined 13C pulse-labelling of 8-year-old beech trees (Fagus sylvatica L.) growing outdoors in a nursery with an anatomical study of the phloem tissue in their stems to examine how drought alters carbon transport and phloem transport capacity. For the six trees under drought, predawn leaf water potential ranged from -0.7 to -2.4 MPa, compared with an average of -0.2 MPa in five control trees with no water stress. We also observed a longer residence time of excess 13C in the foliage and the phloem sap in trees under drought compared with controls. Compared with controls, excess 13C in trunk respiration peaked later in trees under moderate drought conditions and showed no decline even after 4 days under more severe drought conditions. We estimated higher phloem sap viscosity in trees under drought. We also observed much smaller sieve-tube radii in all drought-stressed trees, which led to lower sieve-tube conductivity and lower phloem conductance in the tree stem. We concluded that prolonged drought affected phloem transport capacity through a change in anatomy and that the slowdown of phloem transport under drought likely resulted from a reduced driving force due to lower hydrostatic pressure between the source and sink organs.

Analysis of the age structure of Dahurian larch (Larix gmelinii Rupr.) forests in the western part of the Putorana Plateau (Sukhie Gory Range) and comparison of diachronous topographic maps and satellite images have revealed changes in the altitudinal position of the upper boundary of tree vegetation. the most significant changes occurred on south- and southwest-facing slopes. Correlations of trunk diameters with the phytomass of trees and its fractions have been found. the course of changes in the structure and phytomass of tree stands in the upper timberline over the past centuries has been reconstructed. Forest expansion has been facilitated by the general change in climate conditions (warming) in the study area. This study was supported by the Russian Science Foundation, project no. 17-14-01112.

Interaction of disturbance agents may cause cascading effects in forests. The three most important disturbance agents of Norway spruce (Picea abies) in northern Europe are Heterobasidion root rot, wind and the European spruce bark beetle (Ips typographus). In this study, we present a mechanistic individual agent based model to simulate the dynamics of the bark beetle and integrate it in the simulation framework WINDROT to further study the interactive dynamics of root rot, wind and bark beetles. We carried out model performance analysis concluding that the model is sensitive to the parameters of optimal bark thickness for reproduction. Stand level interaction between wind and bark beetle disturbances was also evaluated against field data. The stand level simulations show the interaction between the disturbance agents; the root rot increases wind disturbance and bark beetles benefit from wind fallen trees. No direct interaction was found in the simulation study between the root rot and bark beetles. Further experimental research and large scale simulation studies are needed to better understand the underlying mechanisms in the interaction between the disturbance agents.

Questions How have tree growth and growth trends varied in structurally complex old-growth forests? What determines this variation and how this differs among species? Location Natural boreal old-growth forests, northern Finland. Methods We sampled trees over 10-cm diameter on 48 randomly located 0.1-ha plots in three landscapes and compiled a data set of 1,565 trees including all three main species, and a full range of sizes and competitive states in northern boreal old-growth forests. We extracted tree ring samples from every tree and used ring widths to reconstruct tree sizes and their neighbourhood for the past 50 years for Pinus sylvestris and Picea abies, and 30 years for Betula pubescens. We used mixed-effects models to study variation in tree growth. In the models, we used annually varying environmental variables, forest and tree structural variables and site variables as predictors. We also measured soil hydraulic properties for a water balance model to specifically include soil moisture deficits as predictors. Results Tree growth rates had increased during the analysis period for all three species, with the highest increases for P. sylvestris and B. pubescens. All three species grew well during warm summers, but P. sylvestris and P. abies growth was also negatively influenced by soil moisture deficits. Competition in these sparse forests had a small influence on growth. However, its influence increased over time. Conclusions The trees showed positive growth trends, but the increasing competition means that such strong positive trends are unlikely to continue over the long term. For year-to-year variability, negative influence of droughts implied that increasing temperatures are not only beneficial to tree growth even in these high-latitude forests. Overall, the findings demonstrate the site- and species-dependent complexity in tree responses to environmental and structural changes we may expect in the future.

Conifer trees, including Norway spruce, are threatened by fungi of the Heterobasidion annosum species complex, which severely affect timber quality and cause economic losses to forest owners. The timely detection of infected trees is complicated, as the pathogen resides within the heartwood and sapwood of infected trees. The presence of the disease and the extent of the wood decay often becomes evident only after tree felling. Fourier-transform infrared (FT-IR) spectroscopy is a potential method for non-destructive sample analysis that may be useful for identifying infected trees in this pathosystem. We performed FT-IR analysis of 18 phloem, 18 xylem, and 18 needle samples from asymptomatic and symptomatic Norway spruce trees. FT-IR spectra from 1066 – 912 cm−1 could be used to distinguish phloem, xylem, and needle tissue extracts. FT-IR spectra collected from xylem and needle extracts could also be used to discriminate between asymptomatic and symptomatic trees using spectral bands from 1657 – 994 cm−1 and 1104 – 994 cm−1, respectively. A partial least squares regression model predicted the concentration of condensed tannins, a defense-related compound, in phloem of asymptomatic and symptomatic trees. This work is the first to show that FT-IR spectroscopy can be used for the identification of Norway spruce trees naturally infected with Heterobasidion spp.

Root and butt rot caused by members of the Heterobasidion annosum species complex is the most economically important disease of conifer trees in boreal forests. Wood decay in the infected trees dramatically decreases their value and causes considerable losses to forest owners. Trees vary in their susceptibility to Heterobasidion infection, but the genetic determinants underlying the variation in the susceptibility are not well-understood. We performed the identification of Norway spruce genes associated with the resistance to Heterobasidion parviporum infection using genome-wide exon-capture approach. Sixty-four clonal Norway spruce lines were phenotyped, and their responses to H. parviporum inoculation were determined by lesion length measurements. Afterwards, the spruce lines were genotyped by targeted resequencing and identification of genetic variants (SNPs). Genome-wide association analysis identified 10 SNPs located within 8 genes as significantly associated with the larger necrotic lesions in response to H. parviporum inoculation. The genetic variants identified in our analysis are potential marker candidates for future screening programs aiming at the differentiation of disease-susceptible and resistant trees.

Climate-related extended outbreaks and range shifts of destructive bark beetle species pose a serious threat to urban boreal forests in North America and Fennoscandia. Recent developments in low-cost remote sensing technologies offer an attractive means for early detection and management of environmental change. They are of great interest to the actors responsible for monitoring and managing forest health. The objective of this investigation was to develop, assess, and compare automated remote sensing procedures based on novel, low-cost hyperspectral imaging technology for the identification of bark beetle infestations at the individual tree level in urban forests. A hyperspectral camera based on a tunable Fabry-Perot interferometer was operated from a small, unmanned airborne vehicle (UAV) platform and a small Cessna-type aircraft platform. This study compared aspects of using UAV datasets with a spatial extent of a few hectares (ha) and a ground sample distance (GSD) of 10-12 cm to the aircraft data covering areas of several km(2) and having a GSD of 50 cm. An empirical assessment of the automated identification of mature Norway spruce (Picea abies L. Karst.) trees suffering from infestation (representing different colonization phases) by the European spruce bark beetle (Ips typographus L.) was carried out in the urban forests of Lahti, a city in southern Finland. Individual spruces were classified as healthy, infested, or dead. For the entire test area, the best aircraft data results for overall accuracy were 79% (Cohen's kappa: 0.54) when using three crown color classes (green as healthy, yellow as infested, and gray as dead). For two color classes (healthy, dead) in the same area, the best overall accuracy was 93% (kappa: 0.77). The finer resolution UAV dataset provided better results, with an overall accuracy of 81% (kappa: 0.70), compared to the aircraft results of 73% (kappa: 0.56) in a smaller sub-area. The results showed that novel, low-cost remote sensing technologies based on individual tree analysis and calibrated remote sensing imagery offer great potential for affordable and timely assessments of the health condition of vulnerable urban forests.

The maternal temperature during seed development can significantly affect seed dormancy, germination and seedling performance. While the response of germination and seedling phenology to maternal temperatures has been well studied for annuals and conifers, very few studies focus on deciduous trees. To understand the responses of seedlings to variation in maternal temperature during seed maturation, we assessed the germination, bud phenology (bud burst, bud set) and height of full sib families in a common garden. We performed three controlled crosses between three different pairs of genotypes of European black poplar (Populus nigra) to achieve full sib families in three experiments in warm (+ 10 degrees C) and cold (control) maternal environments during crossing and seed maturation. Warmer (+ 10 degrees C) maternal temperatures decreased the seed germination success. The seedlings from the warmer maternal environment also displayed later bud burst and earlier bud set, but only in one out of the three crossings (Proven female x Horrues male). Our results indicate that the maternal environment can considerably impact on seed germination and the phenological responses of even two-year old seedlings suggesting the existence of a memory of maternal temperature during seed maturation. The seedlings resulting from the colder maternal environment grew taller than those from the warmer environment during the first, but not second, growing season. Our results further our understanding of the responses of deciduous forest trees to rapid climate change, but more research is needed to better understand the mechanisms behind the observed effects of maternal warming.

Trees play a key role in the global hydrological cycle and measurements performed with the thermal dissipation method (TDM) have been crucial in providing whole-tree water-use estimates. Yet, different data processing to calculate whole-tree water use encapsulates uncertainties that have not been systematically assessed. We quantified uncertainties in conifer sap flux density (F-d) and stand water use caused by commonly applied methods for deriving zero-flow conditions, dampening and sensor calibration. Their contribution has been assessed using a stem segment calibration experiment and 4yr of TDM measurements in Picea abies and Larix decidua growing in contrasting environments. Uncertainties were then projected on TDM data from different conifers across the northern hemisphere. Commonly applied methods mostly underestimated absolute F-d. Lacking a site- and species-specific calibrations reduced our stand water-use measurements by 37% and induced uncertainty in northern hemisphere F-d. Additionally, although the interdaily variability was maintained, disregarding dampening and/or applying zero-flow conditions that ignored night-time water use reduced the correlation between environment and F-d. The presented ensemble of calibration curves and proposed dampening correction, together with the systematic quantification of data-processing uncertainties, provide crucial steps in improving whole-tree water-use estimates across spatial and temporal scales.

Several young damaged Norway spruce stands in eastern and central Finland were observed from 2013 to 2016. The damage included trees with heavy resin flow, necrotic foliage, stem and branch cankers and dead trees. Pest identification resulted in the tortricid moth Cydia pactolana whose occurrence was always associated with the presence of the ascomycete pathogen Neonectria fuckeliana. Both the insect and the disease contributed to the extent of the damage, but it is not possible to say in which order they had attacked the trees. Apparently, changed climate has affected the increased occurrence of both the fungus and the moth. However, the characteristics of the insect-fungus interaction and the factors contributing to the coincidences are unknown. Emerging coexistence or potential symbiosis of the two damaging agents is a serious threat for Norway spruce cultivation. Understanding the biology of this fungus-insect interaction is important for controlling them.

Norway spruce is one of the most important commercial forestry species in Europe, and is commonly infected by the bark beetle-vectored necrotrophic fungus, Endoconidiophora polonica. Spruce trees display a restricted capacity to respond to environmental perturbations, and we hypothesized that water limitation will increase disease severity in this pathosystem. To test this prediction, 737 seedlings were randomized to high (W+) or low (W−) water availability treatment groups, and experimentally inoculated with one of three E. polonica strains or mock-inoculated. Seedling mortality was monitored throughout an annual growing season, and total seedling growth and lesion length indices were measured at the experiment conclusion. Seedling growth was greater in the W+ than W− treatment group, demonstrating limitation due to water availability. For seedlings infected with two of the fungal strains, no differences in disease severity occurred in response to water availability. For the third fungal strain, however, greater disease severity (mortality and lesion lengths) occurred in W− than W+ seedlings. While the co-circulation in nature of multiple E. polonica strains of varying virulence is known, this is the first experimental evidence that water availability can alter strain-specific disease severity.

New Phytol.ISI Document Delivery No.: CM6TZTimes Cited: 0Cited Reference Count: 59Charrier, Guillaume Pramsohler, Manuel Charra-Vaskou, Katline Saudreau, Marc Ameglio, Thierry Neuner, Gilbert Mayr, StefanAnr [i826-b25]; fwf [i826-b25]This project was funded by the French and Austrian research agencies (ANR and FWF), project I826-B25 'Acoufreeze'.Wiley-blackwellHoboken; Ultrasonic acoustic emission analysis enables nondestructive monitoring of damage in dehydrating or freezing plant xylem. We studied acoustic emissions (AE) in freezing stems during ice nucleation and propagation, by combining acoustic and infrared thermography techniques and controlling the ice nucleation point. Ultrasonic activity in freezing samples of Picea abies showed two distinct phases: the first on ice nucleation and propagation (up to 50AEs(-1); reversely proportional to the distance to ice nucleation point), and the second (up to 2.5AEs(-1)) after dissipation of the exothermal heat. Identical patterns were observed in other conifer and angiosperm species. The complex AE patterns are explained by the low water potential of ice at the ice-liquid interface, which induced numerous and strong signals. Ice propagation velocities were estimated via AE (during the first phase) and infrared thermography. Acoustic activity ceased before the second phase probably because the exothermal heating and the volume expansion of ice caused decreasing tensions. Results indicate cavitation events at the ice front leading to AE. Ultrasonic emission analysis enabled new insights into the complex process of xylem freezing and might be used to monitor ice propagation in natura.

Apoplastic events such as monolignol oxidation and lignin polymerization are difficult to study in intact trees. To investigate the role of apoplastic hydrogen peroxide (H2O2) in gymnosperm phenolic metabolism, an extracellular lignin-forming cell culture of Norway spruce (Picea abies) was used as a research model. Scavenging of apoplastic H2O2 by potassium iodide repressed lignin formation, in line with peroxidases activating monolignols for lignin polymerization. Time-course analyses coupled to candidate substrate-product pair network propagation revealed differential accumulation of low-molecular-weight phenolics, including (glycosylated) oligolignols, (glycosylated) flavonoids, and proanthocyanidins, in lignin-forming and H2O2-scavenging cultures and supported that monolignols are oxidatively coupled not only in the cell wall but also in the cytoplasm, where they are coupled to other monolignols and proanthocyanidins. Dilignol glycoconjugates with reduced structures were found in the culture medium, suggesting that cells are able to transport glycosylated dilignols to the apoplast. Transcriptomic analyses revealed that scavenging of apoplastic H2O2 resulted in remodulation of the transcriptome, with reduced carbon flux into the shikimate pathway propagating down to monolignol biosynthesis. Aggregated coexpression network analysis identified candidate enzymes and transcription factors for monolignol oxidation and apoplastic H2O2 production in addition to potential H2O2 receptors. The results presented indicate that the redox state of the apoplast has a profound influence on cellular metabolism.

Climate changes, exemplified by increased temperatures and CO2 concentration, pose a global threat to forest health. Of particular concern are pests and pathogens, with a warming climate altering their distributions and evolutionary capacity, while impairing the ability of some plants to respond to infections. Progress in understanding and mitigating such effects is currently hindered by a lack of empirical research. Norway spruce (Picea abies) is one of the most economically important tree species in northern Europe, and is considered highly vulnerable to changes in climate. It is commonly infected by the fungus Endoconidiophora polonica, and we hypothesized that damage caused to trees will increase under future climate change predictions. To test this hypothesis an in vivo greenhouse experiment was conducted to evaluate the effects of a changed growing environment on E. polonica infected Norway spruce seedlings, comparing ambient conditions to predicted temperatures and CO2 levels in Finland for the years 2030 and 2100. In total, 450 seedlings were randomized amongst the three treatments, with 25 seedlings from each allocated to inoculation with one of five different fungal strains or mock-inoculation. Seedlings were monitored throughout the thermal growing season for mortality, and lesion length and depth indices were measured at the experiment conclusion. Disease severity (mortality and lesions) was consistently greater in fungal-inoculated than mock-inoculated seedlings. However, substantial differences were observed among fungal strains in response to climate scenarios. For example, although overall seedling mortality was highest under the most distant (and severe) climate change expectations, of the two fungal strains with the highest mortality counts (referred to as F4 and F5), one produced greater mortality under the 2030 and 2100 scenarios than ambient conditions, whereas climate scenario had no effect on the other. This study contributes to a limited body of empirical research on the effects of projected climate changes on forestry pathosystems, and is the first to investigate interactions between Norway spruce and E. polonica. The results indicate the potential for future climate changes to alter the impact of forest pathogens with implications for productivity, while highlighting the need for a strain-specific level of understanding of the disease agents.

Aim of study: We aim at (i) developing a reference definition of mixed forests in order to harmonize comparative research in mixed forests and (ii) review the research perspectives in mixed forests.Area of study: The definition is developed in Europe but can be tested worldwide.Material and Methods: Review of existent definitions of mixed forests based and literature review encompassing dynamics, management and economic valuation of mixed forests.Main results: A mixed forest is defined as a forest unit, excluding linear formations, where at least two tree species coexist at any developmental stage, sharing common resources (light, water, and/or soil nutrients). The presence of each of the component species is normally quantified as a proportion of the number of stems or of basal area, although volume, biomass or canopy cover as well as proportions by occupied stand area may be used for specific objectives. A variety of structures and patterns of mixtures can occur, and the interactions between the component species and their relative proportions may change over time.The research perspectives identified are (i) species interactions and responses to hazards, (ii) the concept of maximum density in mixed forests, (iii) conversion of monocultures to mixed-species forest and (iv) economic valuation of ecosystem services provided by mixed forests.Research highlights: The definition is considered a high-level one which encompasses previous attempts to define mixed forests. Current fields of research indicate that gradient studies, experimental design approaches, and model simulations are key topics providing new research opportunities. Aim of study: We aim at (i) developing a reference definition of mixed forests in order to harmonize comparative research in mixed forests and (ii) review the research perspectives in mixed forests.Area of study: The definition is developed in Europe but can be tested worldwide.Material and Methods: Review of existent definitions of mixed forests based and literature review encompassing dynamics, management and economic valuation of mixed forests.Main results: A mixed forest is defined as a forest unit, excluding linear formations, where at least two tree species coexist at any developmental stage, sharing common resources (light, water, and/or soil nutrients). The presence of each of the component species is normally quantified as a proportion of the number of stems or of basal area, although volume, biomass or canopy cover as well as proportions by occupied stand area may be used for specific objectives. A variety of structures and patterns of mixtures can occur, and the interactions between the component species and their relative proportions may change over time.The research perspectives identified are (i) species interactions and responses to hazards, (ii) the concept of maximum density in mixed forests, (iii) conversion of monocultures to mixed-species forest and (iv) economic valuation of ecosystem services provided by mixed forests.Research highlights: The definition is considered a high-level one which encompasses previous attempts to define mixed forests. Current fields of research indicate that gradient studies, experimental design approaches, and model simulations are key topics providing new research opportunities.

International audience; During the 1980s and 1990s, liming was largely used as a tool to protect or restore forest stands located on acid soils from further acidification due to atmospheric deposition. Whereas impacts on soil properties and tree nutrition are well documented, little is known about the long-term temporal impact of liming on radial growth, and on the effect of climate on this response. We therefore analysed time series of radial increment on 128 dominant trees from spruce liming trials located in the Belgian (Gouvy, 1995-2008) and French (La Croix-Scaille, 1981-2008) Ardennes; these stands were limed at ca 40-45 years old. The growth gain associated with liming (GGL, %) was defined by the ratio of the difference in Mean Individual Basal Area Increment between treatments (BAINC limed-BAINC control) corrected for initial differences, to BAINC of the control trees. The effect of climate on GGL was assessed by computing the difference between precipitation (P) and evapotranspiration (PET) over three reference periods of the current growing year (March-June, July-September, March-September). Liming resulted in a two stage growth response pattern. In the first stage (4-8 years after treatment), GGL increased up to a maximum value of 29% in Gouvy and 54% in La Croix-Scaille. In a second stage, GGL decreased linearly with time from application while remaining >0. In La Croix-Scaille, the residuals around this tendency (Y) were positively and linearly related to the difference (P-PET; X) during the current vegetation period (Y= 0.0037X - 0.68; R-2 = 0.36, P = 0.0029), suggesting tree response to liming depended on water availability. Compared to the control, liming increased the expected growth response that is described by the linear trend, without considering the variation due to climate. (C) 2012 Elsevier B.V. All rights reserved.

The quantification of cambial growth over short time periods has been hampered by problems to discern between growth and the swelling and shrinking of a tree stem. This paper presents a model, which separates cambial growth and reversible water-potential induced diurnal changes from simultaneously measured whole stem and xylem radial variations, from field-measured Scots pine trees in Finland. The modelled growth, which includes osmotic concentration changes, was compared with (direct) dendrometer measurements and microcore samples. In addition, the relationship of modelled growth and dendrometer measurements to environmental factors was analysed. The results showed that the water-potential induced changes of tree radius were successfully separated from stem growth. Daily growth predicted by the model exhibited a high correlation with the modelled daily changes of osmotic concentration in phloem, and a temperature dependency in early summer. Late-summer growth saw higher dependency on water availability and temperature. Evaluation of the model against dendrometer measurements showed that the latter masked a true environmental signal in stem growth due to water-potential induced changes. The model provides better understanding of radial growth physiology and offers potential to examine growth dynamics and changes due to osmotic concentration, and how the environment affects growth.

COST Action FP1106 STReESS ; EU Life Programme (LIFE12 ENV/FI/000409 ; Ajut Marie Curie IF fellowship (No 659191) 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.

This study aimed at examining whether plant species of varying resource quality give rise to a fungal- or bacterial-based energy channel in the plants' rhizosphere, when planted in soil, in which the plant species naturally occur. In an 18-month large-scale laboratory mesocosm experiment, two plant species (Holcus lanatus and Lotus corniculatus) producing labile litter and two plant species (Picea abies and Calluna vulgaris) producing recalcitrant litter were placed in the same mesocosm. This allowed the decomposer biota (microbes, protozoa, nematodes, collembolans, and enchytraeid worms) to freely choose their preferred plant rhizosphere. Because a fungal-based energy channel is in theory regarded to retain nutrients better in the soil than a bacterial-based energy channel, water was collected underneath the plant species and analyzed for its nutrient (N and P) content. In general, the number of soil biota groups responding significantly to the plant treatment increased with time. Soil microbes were the first group to react, but the ability of the plants to boost a clearly fungal- or bacterial-based energy channel was generally weak. However, at the end of the study, a higher fungal-to-bacterial biomass ratio was found beneath Calluna than beneath the other plant treatments. Of the secondary consumers, nematodes were the most responsive group, with total number and especially plant parasites being more abundant beneath Lotus than beneath Picea and Calluna indicating a root-based energy channel to persist under the legume. Protozoa, enchytraeid worms and collembolans responded weakly and inconsistently to plant treatment. Liberation of plant-available nutrients was plant-dependent. Towards the end of the study, less nitrate and phosphate leached through the Picea soils than through the control, which indicates a more effective nutrient retention to take place under this plant species. We found that plant species can, in a relatively short time, modify the composition of the soil decomposer community. However, no consistent evidence for the plant-soil systems developing to separated detritus energy channels emerged, we found a clearly separated root energy channel under the legume L. corniculatus.

Water-soluble native O-acetyl galactoglucomannan (GGM) from spruce is a polysaccharide that can be produced in an industrial scale. To develop GGM applications, information is needed on its stability, particularly under acidic conditions. Therefore, acid hydrolysis of spruce GGM was investigated at various pH levels and temperatures. The results allow an estimation of the stability of GGM under food processing conditions and in biological systems. Determination of the average molar mass demonstrated that spruce GGM was stable at pH 1 and 37°C, as well as at pH 3 and 70°C. GGM was hydrolysed at pH 1 and 90°C. GGM oligomers and monomers were detected after degradation. Some of the oligomers contained O-acetyl groups. Monosaccharides were the predominant products in the hydrolysates after treatment at pH 1 and 90°C for 48 h. Pentoses, present in GGM samples as impurities, were released more easily than GGM hexoses. Glucose was more difficult to release than mannose. Traces of 6-deoxy-mannose and levoglucosan were found in the hydrolysates, indicating further degradation of hydrolysed monosaccharides.

Thousands of ectomycorrhizal (ECM) fungal species exist, but estimates of global species richness of ECM fungi differ widely. Many genera have been proposed as being ECM, but ill a number of studies evidence for the hypothesized ECM habit is lacking. Progress in estimating ECM species richness is therefore slow. Ill this paper we have retrieved studies providing evidence for the ECM habit of fungal species and For the identification of the mycobiont(s) ill specific ECM associations, using published and web-based mycorrhiza literature. The identification methods considered are morpho-anatomical, characterization or naturally occurring ECMs, pure Culture synthesis, molecular identification, and isotopic evidence. In addition, phylogenetic information is also considered as a relevant criterion to assess ECM habit. OF 343 fungal genera for which all ECM status has been alleged, about two thirds have Supportive published evidence or ECM Status can be at least hypothesized. For the remaining taxa, Currently no indication exists as for their I-CM nutritional habit, besides field observations or associations with putative hosts. Our survey clearly indicates that current knowledge of ECM fungal diversity, as Supported by experimental evidence, is only partly complete, and that inclusion of many Funga genera in this trophic and ecological category is not verified at this stage. Care must thus be used when compiling lists of ECM and saprotrophic full studies oil the basis of published information only. On the basis of our literature search we conservatively estimate ECM species richness around 7750 species. However, oil the basis of estimates of knowns and unknowns in macromycete diversity, a final estimate or ECM species richness Would likely be between 20000 and 25000.

Summary The aim of this study was to evaluate different silvicultural systems for Douglas-fi r ( Pseudotsuga menziesii (Mirb.) Franco) in the Netherlands in terms of timber production and wind stability over a full rotation. This was done using the forest genetics, ecology, management and wind model (ForGEM-W), which combines a distant dependent tree growth simulator with a mechanical – empirical wind damage module. Six different silvicultural systems were evaluated: normal yield table management, free thinning from above in a monospecies and a mixed stand (50 per cent mixture of beech ( Fagus sylvatica L.)), uneven-aged system, no thinning regime with low initial stand density of Douglas-fi r with and without admixture of beech. Silvicultural systems leading to low height – diameter (h/d) ratios were most successful in avoiding damage. Low h/d ratios were obtained in the system with low stand density and no thinning and in the uneven-aged system by systematically removing trees with the highest ratios during thinning. In particular, the uneven-aged system combined a high timber production with low risk. The use of Douglas-fi r – beech mixtures changed the competition pressure on Douglas-fi r, and thus the h/d ratio and the wind risk. Results from this study indicate that the current trend towards more nature-oriented management could lead to lower wind risks and even to an increase in overall productivity.

Models of the annual development cycle of trees in boreal and temperate regions were reviewed and classified on the basis of their ecophysiological assumptions. In our classification we discern two main categories of tree development: 1) fixed sequence development, which refers to irreversible ontogenetic development leading to visible phenological events such as bud burst or flowering, and 2) fluctuating development, which refers to reversible physiological phenomena such as the dynamics of frost hardiness during winter. As many of the physiological phenomena are partially reversible, we also describe integrated models, which include aspects of both fixed-sequence and fluctuating development. In our classification we further discern simple E-models, where the environmental response stays constant, and more comprehensive ES-models, where the environmental response changes according to the state of development. On the basis of this model classification, we have developed an operational modelling framework, in which we define an explicit state variable and a corresponding rate variable for each attribute of the annual cycle considered. We introduce a unifying notation, which we also use when presenting a selection of previously published models. To illustrate the various developmental phenomena and their modelling, we have carried out model simulations. Finally, we discuss the ecophysiological interpretation of the model variables, methodological aspects of the empirical development and testing of the models, the introduction of new aspects to the modelling, other closely related models, and applications of the models.

Heterobasidion annosum sensu lato is a serious pathogen causing root and stem rot to conifers in the northern hemisphere and rendering the timber defective for sawing and pulping. In this study we applied next-generation sequencing to i) identify transcriptional responses unique to Heterobasidion-inoculated Norway spruce and ii) investigate the H. annosum transcripts to identify putative virulence factors. To address these objectives we wounded or inoculated 30-year-old Norway spruce clones with H. annosumand 454-sequenced the transcriptome of the interaction at 0, 5 and 15 days post inoculation. The 491860 high-quality reads were de novo assembled and the relative expression was analysed. Overall, very few H. annosum transcripts were represented in our dataset. Three delta-12 fatty acid desaturase transcripts and one Clavaminate synthase-like transcript, both associated with virulence in other pathosystems, were found among the significantly induced transcripts. The analysis of the Norway spruce transcriptional responses produced a handful of differentially expressed transcripts. Most of these transcripts originated from genes known to respond to H. annosum. However, three genes that had not previously been reported to respond to H. annosum showed specific induction to inoculation: an oxophytodienoic acid-reductase (OPR), a beta-glucosidaseand a germin-like protein (GLP2) gene. Even in a small data set like ours, five novel highly expressed Norway spruce transcripts without significant alignment to any previously annotated protein in Genbank but present in the P. abies (v1.0) gene catalogue were identified. Their expression pattern suggests a role in defence. Therefore a more complete survey of the transcriptional responses in the interactions between Norway spruce and its major pathogen H. annosumwould probably provide a better understanding of gymnosperm defence than accumulated until now.

International audience; Many forest stands grow on acid and nutrient poor soils. To better understand how they cope with very low mineral resources, we investigated (1) Mg and Ca uptake in relation to depth, and (2) the allocation of these elements from the roots to the canopy, using a multi-isotopic (Mg-26, Ca-44) tracing experiment in a beech stand on a very poor soil. The distribution of the tracers in the soil was taken from van der Heijden et al. (Plant Soil 369:33-45, 2013a, Geoderma 195-196:12-22, 2013b, For Ecol Manag 293:65-78, 2013c). A model simulating Mg, Ca, Mg-26 and Ca-44 uptake was developed and applied to estimate the vertical distribution of Mg and Ca uptake in the soil profile. The vertical distribution of tracers in aboveground biomass was measured from four felled trees 2 years after the application of the tracers. The modeled distribution of root uptake in relation to depth shows differences between Mg and Ca: the main source of Mg uptake is the litter layer (circa. 43 % of total uptake) and the top mineral soil (0-5 cm) for Ca (circa. 42 %). The deeper soil layers (15-60 cm) also contribute to uptake. The study does not show clear evidence that uptake occurs in the very deep soil layers (> 70 cm). The distribution of tracers in the aboveground biomass shows a vertical gradient from the stump to the canopy with no or very small amounts of tracers being observed in the foliage during the 2 years after the application of tracers. This suggests that Mg and Ca transport from roots to leaves along the xylem sap flow is quite slow. As Ca and Mg supply to the trees from deep soil horizons is not evidenced, and tracer transport from roots to the leaves is slow, we suggest that the tree internal pool of Ca and Mg may be more active than previously thought. This pool may act as a buffer when nutrient availability is in shortage.