Winter physiological processes are often overlooked in climate change studies of boreal forests, despite their critical role in determining tree survival and carbon balance. Winter climate change is reshaping the environmental constraints that govern boreal forest regeneration, placing increasing pressure on the resilience of tree seedlings. The present mini-review summarises how changing non-growing season temperatures and snow conditions influence the seasonal survival of boreal tree seedlings and the carbon dynamics of both seedlings and soil, with implications for successful forest regeneration. Snow insulates boreal vegetation from extreme cold, regulates light and UV exposure, and shapes spring hydrology. Alterations in snow depth, structure, duration, and melt timing modify soil temperatures, gas exchange, and the snow-soil-root-microbe system, increasing root stress and mortality through deeper frost and hypoxia, ultimately resulting in impaired spring recovery. Winter remains one of the most uncertain parts of the boreal carbon budget, as soils, microbes, and roots continue emitting CO2 and climate change is altering these fluxes and their carryover effects. Boreal forests rely on tightly regulated annual cycles in which cold acclimation and frost hardiness are essential for winter survival, yet warming can delay acclimation, increase vulnerability to warm spells, and advance spring phenology, thereby raising frost-damage risk for seedlings. The post-planting resilience of seedlings can be strengthened by targeted silvicultural planning, nursery, and planting practices that better prepare seedlings to withstand increasingly variable and challenging winter conditions. More research is needed on how boreal forest tree seedlings physiologically and phenologically adapt to changing winter conditions.

The bilberry (Vaccinium myrtillus L.) is an essential wild berry species in Northern Europe, valued for its sweet taste and chemical compounds. The growing environment, particularly the light and temperature conditions, affects the quality of bilberry fruits. However, studies of their quality in boreal forests are scarce. We analyzed the total phenolics, anthocyanins (ACN), flavonols, iridoids and sugars of bilberry fruits collected from 28 forests in central Finland, to identify the characteristics of forest structure, understorey, and soil that best indicate fruit quality. Forest structure was characterized by canopy cover (leaf area index, LAI), mean age and dominant tree species. The mean (f SD) values of the total phenolics, ACNs, iridoids and sugars in bilberry fruits were 88.7 f 14.2, 78.9 f 13.4, 0.52 f 0.11 and 339.1 f 44.6 mg g- 1 d.w., respectively. Based on linear mixed models, forest structure was the strongest predictor of high phenolics and ACNs, while soil organic layer best explained sugar concentrations. Moreover, daily minimum temperatures correlated with higher sugar levels. A significant interaction between the dominant tree species and LAI indicated that light availability increased phenolic content in bilberry fruits in spruce dominated forests. When comparing different forest stand types, bilberry fruits from drained peatland pine forests exhibited the highest levels of ACNs. High ACNs were associated with high graminoid abundance in the understorey. These findings could direct the harvesting of high-quality bilberry fruits and reveal the intricate relationship between bilberry fruit quality, forest stands and site characteristics in natural ecosystems.

The physical and chemical properties of wild berry fruits change dramatically during development, and the ripe berries host species-specific endophytic communities. However, the development of fungal endophytic communities during berry ripening is unknown. We studied bilberries (Vaccinium myrtillus L.), valuable natural resources in northern Europe and richest sources of phenolic compounds, to characterize dynamics of the fungal communities over fruit developmental stages (raw, veraison, and ripe). Our focus was to examine the changes in the fruit phenolic compounds associated with the fungal community structure using liquid chromatography-mass spectrometry for phenolic compounds and high-throughput sequencing technology targeting the internal transcribed spacer 2 ribosomal DNA region for endophytic fungi. We found that the fungal diversity increased with the ripening stages. The fungal profile changed dramatically through fruit development, and the veraison stage was a transition stage, where the core mycobiome of fruits changed. The fungal community structure and abundance of the most dominant genera in raw and ripe stages, Monilinia and Cladosporium, respectively, were driven by the bilberry phenolic profile. We conclude that sampling time, tissue age, and phenolic compounds play important roles in the development of fruit fungal community. Moreover, phenolic compounds could be the host's strategy to recruit beneficial microbes. The diversity and profile of fungal endophytic communities change dramatically through bilberry fruit development; sampling time, tissue age, and host phenolic compounds shape the fungal communities.

Stinging nettle (Urtica dioica) is an herbaceous perennial plant native to Eurasia, wildly distributed throughout the temperate parts of the world. Although generally considered as a weed due to its fast growth and invasive capacity, stinging nettle is well suited to cultivation and is currently experiencing a revival as a beneficial crop due to its numerous potential applications. This interest reflects in an increasing number of scientific articles related to nettle in the last years. However, reports mostly focus on the aerial parts of the plant. Roots are rich in numerous phytochemicals such as phytosterols, lignans, coumarins, sugars, and lectins. By compiling the most relevant publications, the aim of this review is to gather the current knowledge about nettle roots, such as root system functioning, biochemical composition, and related functional activities. A special emphasis is placed on lectins (or UDA for Urtica dioica agglutinin) due to their functional activities. This review highlights the potential of nettle root as a source of biomolecules. Gaps of knowledge and possible future directions for nettle root research, production, and uses are discussed.

Marginal lands have been proposed to produce non-food crop biomass for energy or green materials. For this purpose, the selection, implementation, and growth optimization of plant species on such lands are key elements to investigate to achieve relevant plant yields. Stinging nettle (Urtica dioica) is a herbaceous perennial that grows spontaneously on contaminated lands and was described as suitable to produce fibers for material applications. Two mercury-contaminated soils from industrial wastelands with different properties (grassland soil and sediment landfill) were used in this study to assess the potential growth of stinging nettle in a greenhouse mesocosm experiment. Two organic amendments were studied for their impact on nettle growth. The solid digestate from organic food wastes significantly doubled plant biomass whereas the compost from green wastes had a lower impact. The highest doses of organic amendments significantly increased the number of fibers, which doubled following digestate application, while reducing leaf Hg concentration. Both amendments significantly improved soil respiration and enzymatic activities linked to the microbial biomass in the soil from the sediment landfill by the end of the experiment. In the context of a phytomanagement scenario, solid digestate would be a preferred amendment resource to improve nettle production on industrial wastelands.

Abstract The aboveground parts of boreal forest trees grow earlier in the growing season, the roots mostly later. The idea was to examine whether root growth followed soil temperature, or whether shoot growth also demanded most resources in the early growing season (soil temperature vs internal sink strengths for resources). The linkage between air and soil temperature was broken by switching the soil temperature. We aimed here to identify the direct effects of different soil temperature patterns on physiology, leaf anatomy and their interactions, and how they relate to the control of the growth dynamics of silver birch (Betula pendula Roth). Sixteen 2-year-old seedlings were grown in a controlled environment for two 14-week simulated growing seasons (GS1, GS2). An 8-week dormancy period interposed the GSs. In GS2, soil temperature treatments were applied: constant 10 °C (Cool), constant 18 °C (Warm), early growing season at 10 °C switched to 18 °C later (Early Cool Late Warm) and 18 °C followed by 10 °C (Early Warm Late Cool) were applied during GS2. The switch from cool to warm enhanced the water status, net photosynthesis, chlorophyll content index, effective yield of photosystem II (ΔF/Fm′) and leaf expansion of the seedlings. Warm treatment increased the stomatal number per leaf. In contrast, soil cooling increased glandular trichomes. This investment in increasing the chemical defense potential may be associated with the decreased growth in cool soil. Non-structural carbohydrates were accumulated in leaves at a low soil temperature showing that growth was more hindered than net photosynthesis. Leaf anatomy differed between the first and second leaf flush of silver birch, which may promote tree fitness in the prevailing growing conditions. The interaction of birch structure and function changes with soil temperature, which can further reflect to ecosystem functioning.

In northern boreal forests the warming winter climate leads to more frequent snowmelt, rain-on-snow events and freeze-thaw cycles. This may be harmful or even lethal for tree seedlings that spend even a half of the year under snow. We conducted a snow cover manipulation experiment in a natural forest to find out how changing snow conditions affect young Scots pine (Pinus sylvestris L.) seedlings. The ice encasement (IE), absence of snow (NoSNOW) and snow compaction (COMP) treatments affected ground level temperature, ground frost and subnivean gas concentrations compared to the ambient snow cover (AMB) and led to the increased physical damage and mortality of seedlings. The expression responses of 28 genes related to circadian clock, aerobic and anaerobic energy metabolism, carbohydrate metabolism and stress protection revealed that seedlings were exposed to different stresses in a complex way depending on the thickness and quality of the snow cover. The IE treatment caused hypoxic stress and probably affected roots which resulted in reduced water uptake in the beginning of the growing season. Without protective snowpack in NoSNOW seedlings suffered from cold and drought stresses. The combination of hypoxic and cold stresses in COMP evoked unique transcriptional responses including oxidative stress. Snow cover manipulation induced changes in the expression of several circadian clock related genes suggested that photoreceptors and the circadian clock system play an essential role in the adaptation of Scots pine seedlings to stresses under different snow conditions. Our findings show that warming winter climate alters snow conditions and consequently causes Scots pine seedlings various abiotic stresses, whose effects extend from overwintering to the following growing season.

ABSTRACT Wild berries are interesting research subjects due to their rich sources of health-beneficial phenolic compounds. However, the internal microbial communities, endophytes, associated with the wild berry fruits are currently unknown. Endophytes are bacteria or fungi inhabiting inside plant tissues, and their functions vary depending on the host species and environmental parameters. The present study aimed to examine community composition of fungal and bacterial endophytes in fruits of three wild berry species (bilberry Vaccinium myrtillus L., lingonberry Vaccinium vitis-idaea L. and crowberry Empetrum nigrum L.) and the effects of host plant species and their growth sites on shaping the endophytic communities. We found that the endophytic community structures differed between the berry species, and fungi were predominant over bacteria in the total endophytic taxa. We identified previously unknown endophytic fungal taxa including Angustimassarina, Dothidea, Fellozyma, Pseudohyphozyma, Hannaella coprosmae and Oberwinklerozyma straminea. A role of soluble phenolic compounds, the intracellular components in wild berry fruits, in shaping the endophytic communities is proposed. Overall, our study demonstrates that each berry species harbors a unique endophytic community of microbes.

Future climate scenarios for the boreal zone project increasing temperatures and precipitation, as well as extreme weather events such as heavy rain during the growing season. This can result in more frequent short-term waterlogging (WL) leading to unfavorable conditions for tree roots. In addition, it is decisive whether short-term WL periods during the growing season occur continuously or periodically. We assessed the effects of short-termed WL on 4-year-old Scots pine (Pinus sylvestris L.) saplings after shoot elongation started. Waterlogging (WL) lasted either continuously for 2.5 weeks (ContWL) or noncontinuously for 5 weeks, consisting of three repeated 1-week-interval WL periods (IntWL). Both treatments resulted in the same duration of soil anoxia. We studied soil gases, root and shoot growth and physiology, and root survival probability and longevity during the experiment. In the final harvest, we determined shoot and root biomass and hydraulic conductance and electrical impedance spectra of the root systems. Soil CO2 and CH4 concentrations increased immediately after WL onset and O2 decreased until anoxia. Waterlogging decreased fine root survival probability, but there was no difference between WL treatments. Shoot growth suffered more from ContWL and root growth more from IntWL. Needle concentrations of pinitol increased in the WL saplings, indicating stress. No WL effects were observed in photosynthesis and chlorophyll fluorescence. Increased starch concentration in needles by WL may be due to damaged roots and thus a missing belowground sink. Electrical impedance indicated suffering of WL saplings, although root hydraulic conductance did not differ between the treatments. Oxidative stress of short-term and interval WL can have long-lasting effects on shoot and root growth and the physiology of Scots pine. We conclude that even short-term WL during the growing season is a stress factor, which will probably increase in the future and can affect carbon allocation and dynamics in boreal forests.

In boreal forests, snow typically covers the forest floor and insulates soil from low and fluctuating air temperature for a large part of the year. Snowpack regimes are currently changing in response to climate warming, and it is uncertain how changing subnivean conditions (i.e. those under the snow cover) will impact soil processes. Here, we experimentally tested in a northern boreal forest how three various types of snow cover changes 1) ice encasement, 2) the absence of snow, and 3) snow compaction influence wintertime soil gas concentrations, growing season microbial respiration and biomass, extracellular enzyme activities (EEAs), and dissolved organic carbon (DOC) and nitrogen (N). The consequences varied drastically among experimental treatments. An initially strong increase in wintertime soil CO2 accumulation in response to ice encasement weakened during the course of the 3-year experiment, when at the same time, growing season phenol oxidase decreased by 13% and β-glucosidase activity increased by 25%. The absence of snow enhanced β-glucosidase activity by 47% and, by the third year of experiment, increased inorganic soil N concentration. Snow compaction, in turn, enhanced phenol oxidase activity by 27% together with soil microbial respiration and biomass carbon. These findings suggest that consequences of snowpack regime change on the boreal forest soil carbon and nutrient cycling may vary from positive to negative depending on the type of change in the snow cover that becomes dominant.

Our objective was to study how changes in the light spectra affect growth; the concentration of carbohydrate, chlorophyll, carotenoid, terpene, alkaloid, and phenolic compounds; and the emissions of BVOC (biogenic volatile organic compound) of Norway spruce (Picea abies (L.) Karst.) seedlings. This study was conducted during the growth of the third needle generation in plant growth chambers. The plants were exposed to two light spectra with equal photon flux densities but different proportions of blue light (400–500 nm) produced by LED (light-emitting diode) lamps: (i) control (white light + 12% blue light) and (ii) increased blue light (+B) (white light + 45% blue light). The +B treatment increased the concentrations of total flavonoids and acetophenones in needles. The major changes in the phenolic profile were an accumulation of astragalin derivatives and the aglycone of picein. The +B treatment also decreased concentrations of the main alkaloid compound, epidihydropinidine, and its precursor, 2-methyl-6-propyl-1,6-piperideine; the emission rates of limonene, myrcene, and total monoterpenes; and the concentrations of a few terpenoid compounds, mainly in stems. Growth as well as the carbohydrates and pigments present in the needles were not affected. The results suggest that supplemental blue light shifts carbon allocation between secondary metabolism routes — from alkaloid and terpenoid synthesis to flavonoid and acetophenone synthesis. The changes may affect herbivory and abiotic stress tolerance of Norway spruce.

Air temperatures and precipitation are predicted to increase in the future, especially at high latitudes and particularly so during winter. In contrast to air temperatures, changes in soil temperatures are more difficult to predict, as the fate of the insulating snow cover is crucial in this respect. Soil conditions can also be affected by rain-on-snow events and warm spells during winter, resulting in freeze-thaw cycles, compacted snow, ice encasement and local flooding. These adverse conditions during winter could counteract the otherwise positive effects of climate change on forest growth and productivity. For studying the effects of different winter and snow conditions on young Downy birch (Betula pubescens Ehrh.) seedlings, we carried out a laboratory experiment with birch seedlings subjected to four different winter scenarios: snow covering the seedlings (SNOW), compressed snow and ice encasement (ICE), flooded and frozen soil (FLOOD) and no snow at all (NO SNOW). After the winter treatments we simulated a spring and early summer period of 9.5 weeks, and monitored the growth by measuring shoot and root biomass of the seedlings, and starch and soluble sugar concentrations. We also assessed the stress experienced by the seedlings by measuring leaf chlorophyll fluorescence and gas exchange. Although no difference in mortality was observed between the treatments, the seedlings in the SNOW and ICE treatments had significantly higher shoot and root biomass compared with those in the FLOOD and NO SNOW treatments. We found higher starch concentrations in roots of the seedlings in the SNOW and ICE treatments, compared with those in the FLOOD and NO SNOW treatments, although photosynthesis did not differ. Our results suggest a malfunction of carbohydrate distribution in the seedlings of the FLOOD and NO SNOW treatments, probably resulting from decreased sinks. The results underline the importance of an insulating and protecting snow cover for small tree seedlings, and that future winters with changed snow pattern might affect the growth of tree seedlings and thus possibly species composition and forest productivity.

Due to a difference in plant resource allocation to reproduction, the males of dioecious plants may be more growth-orientated, whereas females may allocate more resources for synthesizing secondary compounds. This mechanism is considered to cause gender-specific differences in the plant responses to the loss of plant biomass. Here, we tested gender dimorphism in the responses of common juniper (Juniperus communis) to shoot cutting in four juniper populations located in northern boreal forests in Finland. We collected shoots from uncut junipers and from junipers subjected to shoot cutting in the previous year, and analyzed them for their shoot growth as well as phenolic and terpenoid concentrations. There were no differences in foliar phenolic or terpenoid concentrations between the males and the females. Shoot cutting increased phenolic but not terpenoid concentrations, similarly, in both males and females. Our study reveals that the nature of gender dimorphism may differ among species and locations, which should be considered in theories on plant gender dimorphism. Given the similar phenolic and terpene concentrations in both genders, the different sexes in the northern juniper populations might experience equal levels of herbivory. This lack of gender dimorphism in biotic interactions could result from the high need of plant secondary metabolites (PSM) against abiotic stresses, which is typical for juniper at high latitudes.

The northern regions are experiencing considerable changes in winter climate leading to more frequent warm periods, rain-on-snow events and reduced snow pack diminishing the insulation properties of snow cover and increasing soil frost and freeze-thaw cycles. In this study, we investigated how the lack of snow cover, formation of ice encasement and snow compaction affect the size, structure and activities of soil bacterial and fungal communities. Contrary to our hypotheses, snow manipulation treatments over one winter had limited influence on microbial community structure, bacterial or fungal copy numbers or enzyme activities. However, microbial community structure and activities shifted seasonally among soils sampled before snow melt, in early and late growing season and seemed driven by substrate availability. Bacterial and fungal communities were dominated by stress-resistant taxa such as the orders Acidobacteriales, Chaetothyriales and Helotiales that are likely adapted to adverse winter conditions. This study indicated that microbial communities in acidic northern boreal forest soil may be insensitive to direct effects of changing snow cover. However, in long term, the detrimental effects of increased ice and frost to plant roots may alter plant derived carbon and nutrient pools to the soil likely leading to stronger microbial responses.

Future climate scenarios predict increased air temperatures and precipitation, particularly at high latitudes, and especially so during winter. Soil temperatures, however, are more difficult to predict, since they depend strongly on the fate of the insulating snow cover. 'Rain-on-snow' events and warm spells during winter can lead to thaw-freeze cycles, compacted snow and ice encasement, as well as local flooding. These adverse conditions could counteract the otherwise positive effects of climatic changes on forest seedling growth. In order to study the effects of different winter and snow conditions on young Scots pine (Pinus sylvestris L.) seedlings, we conducted a laboratory experiment in which 80 1-year-old Scots pine seedlings were distributed between four winter treatments in dasotrons: ambient snow cover (SNOW), compressed snow and ice encasement (ICE), flooded and frozen soil (FLOOD) and no snow (NO SNOW). During the winter treatment period and a 1.5-month simulated spring/early summer phase, we monitored the needle, stem and root biomass of the seedlings, and determined their starch and soluble sugar concentrations. In addition, we assessed the stress experienced by the seedlings by measuring chlorophyll fluorescence, electric impedance and photosynthesis of the previous-year needles. Compared with the SNOW treatment, carbohydrate concentrations were lower in the FLOOD and NO SNOW treatments where the seedlings had almost died before the end of the experiment, presumably due to frost desiccation of aboveground parts during the winter treatments. The seedlings of the ICE treatment showed dead needles and stems only above the snow and ice cover. The results emphasize the importance of an insulating and protecting snow cover for small forest tree seedlings, and that future winters with changed snow patterns might affect the survival of tree seedlings and thus forest productivity.

At high latitudes, the climate has warmed at twice the rate of the global average with most changes observed in autumn, winter and spring. Increasing winter temperatures and wide temperature fluctuations are leading to more frequent rain-on-snow events and freeze-thaw cycles causing snow compaction and formation of ice layers in the snowpack, thus creating ice encasement (IE). By decreasing the snowpack insulation capacity and restricting soil-atmosphere gas exchange, modification of the snow properties may lead to colder soil but also to hypoxia and accumulation of trace gases in the subnivean environment. To test the effects of these overwintering conditions changes on plant winter survival and growth, we established a snow manipulation experiment in a coniferous forest in Northern Finland with Norway spruce and Scots pine seedlings. In addition to ambient conditions and prevention of IE, we applied three snow manipulation levels: IE created by artificial rain-on-snow events, snow compaction and complete snow removal. Snow removal led to deeper soil frost during winter, but no clear effect of IE or snow compaction done in early winter was observed on soil temperature. Hypoxia and accumulation of CO2 were highest in the IE plots but, more importantly, the duration of CO2 concentration above 5% was 17 days in IE plots compared to 0 days in ambient plots. IE was the most damaging winter condition for both species, decreasing the proportion of healthy seedlings by 47% for spruce and 76% for pine compared to ambient conditions. Seedlings in all three treatments tended to grow less than seedlings in ambient conditions but only IE had a significant effect on spruce growth. Our results demonstrate a negative impact of winter climate change on boreal forest regeneration and productivity. Changing snow conditions may thus partially mitigate the positive effect of increasing growing season temperatures on boreal forest productivity.

Dwarf birch ( Betula nana L.), a dominant deciduous dwarf shrub in many tundra ecosystems, is predicted to increase substantially in abundance due to climate warming. Potential warming-induced changes in the concentrations of phenolic compounds in B. nana leaves could influence the susceptibility of B. nana to environmental stresses; however, only a few studies have investigated the effects of climate warming on the phenolic defence in B. nana . We analysed the responses of phenolic concentrations and amounts in B. nana leaves to factorial treatments of warming and fertilization for 2 years in a subarctic tundra heath. Warming induced a strong decrease in total phenolics, including both flavonols (i.e. quercetin and myricetin derivatives, important defence compounds against oxidative stress) and hydrolysable tannins (HTs, important defence compounds against herbivory). Fertilization exerted weaker effects on phenolic concentrations while significantly increasing the leaf area. Our data indicate that climate warming may decrease the level of phenolic defence in B. nana leaves. Given the important role of these compounds in the plant defence, this phenomenon could potentially increase the susceptibility of B. nana to biotic stresses such as herbivory.