Urban forest ecosystems face increasing abiotic stress due to climate change and anthropogenic pressure, requiring robust indicators of plant adaptation. Hedera helix L., a widespread liana with shoot dimorphism, plays a key role in temperate urban woodlands but remains understudied in terms of multilevel responses to environmental gradients. We assessed vegetative and generative shoots with healthy or damaged leaves across heterogeneous urban forest sites to identify functional responses to volumetric water content (VWC), temperature (t), electrical conductivity (ES), and daily light integral (DLI). VWC and DLI emerged as the key drivers of trait variability. The multilevel adaptive trait responses demonstrated the ecological flexibility of a dominant urban liana. We developed a novel Integrative Ecological Index (IEI) based on normalized trait sub-indices, which captures multilevel plant responses to environmental stress and enables quantitative assessment of urban habitat conditions. IEI was highest in healthy vegetative shoots and lowest in damaged generative shoots. Our findings demonstrate the ecological plasticity of H. helix and its potential as a sensitive bioindicator. This interdisciplinary study provides a scalable trait-based tool to evaluate adaptive capacity and microhabitat conditions in heterogeneous urban environments.

The resource strategy of seedlings is an important key for understanding the adaptation of trees at this ontogenetic phase to abiotic changes. In this study, we tried to determine the patterns of response of functional traits of shade-tolerant and shade-intolerant species along natural environmental light gradients, using an entire-level, trait-based approach by direct and indirect methods in the Arboretum at Kórnik (Poland). Differences between the two species were found for some variables. Analysis of phenotypic plasticity indices of leaf, stem and root traits of seedlings had high values ​​for both species. The values of plasticity indices of A. platanoides root traits were lower compared to the corresponding traits for Q. robur. Relationships between measures obtained from individual-level trait data were stronger than relationships with measures obtained from community-level trait data. The data obtained from the direct method revealed the closest relationships between functional traits of seedlings and light changes at the individual level trait data for both species. Correlation links between LAI and leaf (LMA, SLA) and stem (SSL, SMR) traits were less tight for Q. robur compared to A. platanoides. The indirect phytoindicative method revealed relationships between LC and LMA, and S/R of seedlings based on community-level trait data. Close relationships between LC and LMF, and SLA were not established in contrast to LAI and DIFN. Internal (closest) and external links between the leaf, stem, and root functional traits at the community-level trait data were established.

This study describes the seasonal changes in cell-to-cell transport in three selected angiosperm tree species, Acer pseudoplatanus (maple), Fraxinus excelsior (ash), and Populus tremula × tremuloides (poplar), with an emphasis on the living wood component, xylem parenchyma cells (XPCs). We performed anatomical studies, dye loading through the vascular system, measurements of non-structural carbohydrate content, immunocytochemistry, inhibitory assays and quantitative real-time PCR to analyse the transport mechanisms and seasonal variations in wood. The abundance of membrane dye in wood varied seasonally along with seasonally changing tree phenology, cambial activity, and non-structural carbohydrate content. Moreover, dyes internalized in vessel-associated cells and 'trapped' in the endomembrane system are transported farther between other XPCs via plasmodesmata. Finally, various transport mechanisms based on clathrin-mediated and clathrin-independent endocytosis, and membrane transporters, operate in wood, and their involvement is species and/or season dependent. Our study highlights the importance of XPCs in seasonally changing cell-to-cell transport in both ring-porous (ash) and diffuse-porous (maple, poplar) tree species, and demonstrates the involvement of both endocytosis and plasmodesmata in intercellular communication in angiosperm wood.

Physical and chemical defenses of seeds are essential for plant survival and reproduction. By protecting their seeds from herbivores and other threats, plants can ensure that their offspring have a bet­ter chance of surviving and growing into adult plants. Plant seed-size variation can affect their ability to allocate defense resources and nutrients. Smaller seeds tend to contain fewer resources and are likely to be more constrained in resource allocation to defense or to nutrient supply for the germinating seedling than larger seeds. In this study, we hypothesized that (1) the trade-off between physical and chemical defenses and reserve materials (in this study TNC content) in seeds is related to the allocation of those resources to seeds, 2) the direction of allocation of these resources is mediated by seed size. Based on seeds of 77 woody plant species collected in the Kórnik Arboretum located in western Poland, we measured several physical and chemical seed traits: seed mass, total phenols, tannin content, crude fiber, lignin content, and total non-structural carbohydrates. We revealed that chemical defense (phenol content) and total non-structural carbohydrates increased with seed mass, whereas physical defense (fiber content) had the opposite effect. We found that the trade-off between physical and chemical defense in plants was often mediated by seed size.

Climate change, particularly droughts and heat waves, significantly impacts global photosynthesis and forest ecosystem sustainability. To understand how trees respond to and recover from hydrological stress, we investigated the combined effects of soil moisture and atmospheric vapour pressure deficit (VPD) on seedlings of the two major European broadleaved tree species Fagus sylvatica (FS) and Quercus robur (QR). The experiment was conducted under natural forest gap conditions, while soil water availability was strictly manipulated. We monitored gas exchange (net photosynthesis, stomatal conductance and transpiration rates), nonstructural carbohydrates (NSC) concentration in roots and stomatal morphometry (size and density) during a drought period and recovery. Our comparative empirical study allowed us to distinguish and quantify the effects of soil drought and VPD on stomatal behavior, going beyond theoretical models. We found that QR conserved water more conservatively than FS by reducing transpiration and regulating stomatal conductance under drought. FS maintained higher stomatal conductance and transpiration at elevated VPD until soil moisture became critically low. QR showed higher intrinsic water use efficiency than FS. Stomata density and size also likely played a role in photosynthetic rate and speed of recovery, especially since QR with its seasonal adjustments in stomatal traits (smaller, more numerous stomata in summer leaves) responded and recovered faster compared to FS. Our focal species showed different responses in NSC content under drought stress and recovery, suggesting possible different evolutionary pathways in coping with stress. QR mobilized soluble sugars, while FS relied on starch mobilization to resist drought. Although our focal species often co-occur in mixed forests, our study showed that they have evolved different physiological, morphological and biochemical strategies to cope with drought stress. This suggests that ongoing climate change may alter their competitive ability and adaptive potential in favor of one of the species studied.

AimWarm deserts are characterised by water shortages and high temperature extremes. A commonly reported adaptive strategy in such environments is maximisation of photosynthetic capacity, which allows plants to achieve positive carbon budgets by taking advantage of short periods of water availability and non-inhibitory temperatures. Considering the well-supported interspecific covariation between photosynthetic capacity and leaf N concentration, we tested the hypothesis that environmental aridity is related to an elevated leaf nitrogen content.Location53 locations in the transitional zone spanning the Mediterranean and the Sahara Desert in Morocco. The mean maximal temperature (Tmax) within the area varied between 35.7 degrees C and 43.5 degrees C, and the mean annual precipitation (MAP) was between 12 and 246 mm.Taxon225 vascular species representative of local vegetation.MethodsLeaf samples were collected along a regional aridity gradient and preserved in herbarium presses. The leaf mass per area (LMA) and N concentrations expressed on leaf mass (Nmass) and area (Narea) basis were determined. We also obtained LMA and Nmass values for 6711 species from a worldwide database for comparative analysis.ResultsSignificant increases in mean LMA, Nmass and Narea accompanied the increase in Tmax and the decrease in MAP in woody species and in non-graminoid herbs, but not in graminoids. Considering the overall aridity of our sampling area, we compared the Nmass values of Moroccan plants with those from a worldwide database. We found that at a common LMA, the Moroccan plants showed on average elevated Nmass relative to global values.Main ConclusionsThese two lines of evidence: regional gradient and global comparison confirm that hot deserts select for high leaf N content. This result suggests the direction of natural selection that will accompany future climate warming and habitat aridification.

(1) The focus on floral functional traits and their variability has been significantly lower than when compared to other plant organs. Here, we focused on the variability of four novel floral chemical traits. We aimed to assess the level of interspecific variability of total non-structural carbohydrates (TNC) and total phenolic compounds (TPh) in the flowers of woody species. (2) We collected data on 70 species of temperate trees and shrubs. We also assessed the intraspecific level of variability by collecting flowers from the high-light and low-light parts of the crown. (3) We found a phylogenetic signal in the variability of starch and TPh. We did not observe statistically significant differences or biologically significant trends between the high-light and low-light parts of the crown. We detected a low impact of light availability on the intraspecimen variability of the TNC and TPh contents. (4) Low intraspecimen variability allows for a more reliable extrapolation of measurements in cases of interspecific comparisons and can be used to better describe the reproductive strategies of different woody species.

Trees regenerating in the understory respond to increased availability of light caused by gap formation by undergoing a range of morphological and physiological adjustments. These adjustments include the production of thick, sun-type leaves containing thicker mesophyll and longer palisade cells than in shade-type leaves. We asked whether in the shade-regenerating tree Acer pseudoplatanus , the increase in leaf thickness and expansion of leaf tissues are possible also in leaves that are already fully formed, a response reported so far only for a handful of species. We acclimated potted seedlings to eight levels (from 1 to 100%) of solar irradiance and, in late summer, transferred a subset of them to full sunlight. Within 30 days, the pre-shaded leaves increased leaf mass per area and became thicker mostly due to the elongation of palisade cells, except for the most shaded individuals which suffered irreversible photo-oxidative damage. This anatomical acclimation was accompanied by a transient decline in photosynthetic efficiency of PSII (F v /F M ), the magnitude of which was related to the degree of pre-shading. The F v /F M recovered substantially within the re-acclimation period. However, leaves of transferred plants were shed earlier in the fall, indicating that the acclimation was not fully effective. These results show that A. pseudoplatanus is one of the few known species in which mature leaves may re-acclimate anatomically to increased irradiance. This may be an important mechanism enhancing utilization of gaps created during the growing season.

This dataset consists of combined field transect data and data from a provenance trial. Geographic coordinates and selected Worldclim climatic parameters (mean annual temperature, mean temperature of the warmest quarter and mean annual precipitation) for each sampling site or the original provenance site.are given. For each sampling site we give mean needle length, width, and stomatal traits for each needle surface (number of stomatal rows and stomatal density) as well as the projected stomatal density..

Harsh environmental conditions affect both leaf structure and root traits. However, shoot growth in high‐latitude systems is predominately under photoperiod control while root growth may occur for as long as thermal conditions are favorable. The different sensitivities of these organs may alter functional relationships above‐ and belowground along environmental gradients. We examined the relationship between absorptive root and foliar traits of Scots pine trees growing in situ along a temperate‐boreal transect and in trees grown in a long‐term common garden at a temperate latitude. We related changes in foliar nitrogen, phosphorus, specific leaf area, needle mass and 13C signatures to geographic trends in absorptive root biomass to better understand patterns of altered tree nutrition and water balance. Increased allocation to absorptive fine roots was associated with greater uptake of soil nutrients and subsequently higher needle nutrient contents in the northern provenances compared with more southern provenances when grown together in a common garden setting. In contrast, the leaf δ13C in northern and southern provenances were similar within the common garden suggesting that higher absorptive root biomass fractions could not adequately increase water supply in warmer climates. These results highlight the importance of allocation within the fine‐root system and its impacts on needle nutrition while also suggesting increasing stomatal limitation of photosynthesis in the context of anticipated climatic changes.

Regenerating oak seedlings by different methods impacts taproot architecture, root-to-shoot allometry and, potentially, soil water use and plant growth under water limitations. In the present study, Quercus robur (L.) seedlings regenerated by four different methods - sowing (acorn-sown seedlings), sowing followed by shoot clipping (coppiced), sowing followed by root-pruning (root-pruned), and container planting with the root plug intact (containerized) - were subjected to four water-shortage regimes - full watering and 75, 50, and 25% of full watering. These treatments were used to test the hypotheses that root morphology, anatomy and biochemistry varied among regeneration methods, and, thus, differentially affected plant responses to water-shortages. For most traits, the response to water shortage of acorn-sown and containerized seedlings was similar, and opposite to that of coppiced and root-pruned seedlings. Water shortage did not change tree allocation to total root biomass among regeneration methods but altered the partitioning between absorptive and structural roots within the root system. Acorn-sown and containerized seedlings produced more pioneer roots, and coppiced and root-pruned seedlings more highly-branched fine roots with greater specific root length and specific root surface area under greater water shortage. Thus, acorn-sown and containerized oaks appeared to be primed for water foraging, and coppiced and root-pruned oaks for water absorption efficiency in response to water shortage. Water shortage caused a reduction in nonstructural carbohydrate concentrations in fine roots of acorn-sown and containerized seedlings and in the taproot of coppiced and root-pruned seedlings. Results generally indicated that seedling response to water limitation depended on the regeneration method, highlighting the importance of root-to-shoot allometry, taproot development, and carbon reserve mobilization for plant functioning under water shortage. Acorn-sown and containerized oaks seedlings might be more suitable for forest regeneration in sites characterized by severe water shortage periods. The reduction of taproot starch reserves under strong water shortage may render root-pruned oak trees more susceptible to topsoil water deficit than acorn-sown trees. This work contributes to understand drought acclimation and to identify the most adequate forestry regeneration practices to adapt forest ecosystems to ongoing climatic changes.

The commonly observed negative relationship between stomatal density (SD) and atmospheric CO2 has led to SD being proposed as an indicator of atmospheric CO2 concentration. The use of SD as a proxy for CO2 , however, has been hampered by an insufficient understanding of the intraspecific variation of this trait. We hypothesized that SD in Pinus sylvestris, a widely distributed conifer, varies geographically and that this variation is determined by major climatic variables. By sampling needles from naturally growing trees along a latitudinal range of 32.25°, equivalent to 13.7°C gradient of mean annual temperature (MAT) across Europe, we found that SD decreased from the warmest southern sites to the coldest sites in the north at a rate of 4 stomata per mm2 for each 1°C, with MAT explaining 44% of the variation. Additionally, samples from a provenance trial exhibited a positive relationship between SD and the MAT of the original localities, suggesting that high SD is an adaptation to warm temperature. Our study revealed one of the strongest intraspecific relationships between SD and climate in any woody species, supporting the utility of SD as a temperature, rather than direct CO2 , proxy. In addition, our results predict the response of SD to climate warming.

Key message Juniperus communis males are better adapted than females to changing, seasonal environmental conditions due to their higher photosynthetic capacity and the higher concentration of photosynthetic pigments in their needles. Males cope with ROS more efficiently than females having greater carotenoids concentration in needles. Abstract In dioecious woody plants, females often exhibit greater reproductive effort than male plants and as a result, they can be more vulnerable to different stressors. We hypothesized that female plants of J. communis L. could have a lower photochemical capacity and a higher level of antioxidant enzyme activity and that these differences between males and females would be more pronounced under conditions where nutrient availability is limited. We also assume that additional stressors connected with different seasons would increase those differences. Male and female plants of J. communis growing in fertilized or non-fertilized soils were used to test this hypothesis. The effect of fertilization and sex on photochemical parameters derived from chlorophyll a fluorescence light curves, and on the concentrations of photosynthetic pigments in needles, was determined in different seasons within 2 years. To assess the tolerance of male and female plants to the nutrient deficit, antioxidant enzyme activity, and the level of reactive oxygen species (ROS) were determined. Results revealed sex-related differences in photochemical parameters, level of antioxidant enzyme activity, H 2 O 2 levels, the concentration of photosynthetic pigments, and in the leaf mass-to-area ratio. This indicates that J. communis males could be better adapted than females to changing, seasonal environmental conditions due to their higher photosynthetic capacity, as reflected by their higher ETR max , and a higher concentration of photosynthetic pigments in their needles. The sex-related differences concerning photosynthetic capacity and stress response found in our study are constitutive traits of each sex and are genetically based as they occurred independently of fertilization.

We hypothesized that female and male individuals of the dioecious tree species, Juniperus communis, exhibit different strategies of resource allocation when growing under stress conditions. To test this hypothesis, we performed a two-year pot experiment on plants exposed to different levels of nutrient availability. Analysis of the plants revealed a higher concentration of carbohydrates, carbon, and phenolic compounds in needles of female plants, indicating that females allocate more resources to storage and defense than males. This difference was independent of nutrient availability. Differences in carbohydrates levels between the sexes were most often significant in June, during the most intensive phase of vegetative growth in both sexes, but could also be attributed to female resources investment in cone development. A higher level of nitrogen and other macroelements was observed in males than in females, which may have been connected to the accumulation of resources (nitrogen) for pollen grain production in males or greater allocation of these elements to seeds and cones in females. The interaction between sex and soil fertilization for the C:N ratio may also indicate sex-specific patterns of resource allocation and utilization, which is impacted by their availability during specific periods of J. communis annual life cycle.

Climate change has likely altered high-latitude forests globally, but direct evidence remains rare. Here we show that throughout a ≈1000-km transect in Scots pine (Pinus sylvestris L.) forests in Sweden, mature trees in ≈2015 had longer needles with shorter lifetimes than did trees in ≈1915. These century-scale shifts in needle traits were detected by sampling needles at 74 sites from 2012 to 2017 along the same transect where needle traits had been assessed at 57 sites in 1914-1915. Climate warming of ≈1 °C all along the transect in the past century has driven this temporal shift in foliage traits known to be physiologically critical to growth and carbon cycling processes. These century-scale changes in Scandinavian Scots pine forests represent a fingerprint of climate change on a fundamental biological element, the leaf, with repercussions for productivity and sensitivity to future climate, which are likely to be mirrored by similar changes for evergreen conifers across the boreal biome.

The correlations of phenotypic traits with environmental drivers suggest that variability of these traits is a result of natural selection, especially if such trait correlations are based on genetic variability. We hypothesized that in situ correlations of structural needle traits of Scots pine (Pinus sylvestris L) with minimal winter temperature (Tmin) reported previously from a temperate/boreal transect would be conserved when plants are cultivated under common conditions. We tested this hypothesis by analyzing needles from two common gardens located in the temperate zone, one including adult trees and the other juvenile seedlings. The majority of adult needle traits for which correlations with Tmin were found in the field turned out to be under environmental influence. In contrast, the majority of traits studied in juvenile needles were correlated with the original Tmin suggesting the role of past natural selection in shaping their variability. Juvenile needles thus appeared to be inherently less plastic than adult needles, perhaps reflecting the stronger selective pressure acting during juvenile, as compared with adult, ontogenetic stage. Genetically based cold-climate adaptation in either juvenile or adult needles, or both, involved an increase in leaf mass per area and leaf density, decrease in needle length, reduction in the amount of xylem and phloem, increase in thickness of epidermis, decrease in tracheid diameter and increase in tracheid density, and increase in diameter and volume fraction of resin ducts. We also show that at least some traits, such as transverse xylem and phloem areas and number of fibers, scale with needle length, suggesting that climate-related trait variation may also be mediated by changes in needle length. Moreover, slopes of these allometric relationships may themselves be plastically modified. The phenotypic syndrome typical of needles from cold environments may thus be under environmental, genetic and allometric control.

Key messageIn some lianas the use of trellis modifies within-plant biomass allocation and stimulates growth, however in other species trellis use may negatively affect growth indicating a cost associated with climbing.AbstractIn order to improve access to light, lianas use other plants as climbing trellises. Whereas in shaded lianas climbing a trellis may enhance biomass gain by increasing leaf-level irradiance, we suspected that trellis use may also stimulate growth by triggering modifications in whole-plant allocation of biomass and nitrogen. We evaluated responses to trellis in three temperate lianas: Hedera helix, Celastrus orbiculatus and Wisteria floribunda. Lianas were grown outdoors in 120-L barrels and with ample space, with and without trellises. Biomass and nitrogen accumulation, distribution patterns and total stem length were determined after three and four seasons. Responses were adjusted for individual plant biomass. Liana responses to trellises were species-specific. In W. floribunda trellis use enhanced growth, consistently with the increased leaf mass fraction and leaf area ratio while biomass distribution to roots was reduced. Biomass distribution to stems and total stem length were increased by trellis use. In contrast, growth and biomass distribution in C. orbiculatus were not altered by trellis use. In H. helix growth was reduced in plants climbing on trellises in comparison with plants creeping on the ground, but biomass distribution pattern was not altered. Moreover, supported H. helix plants accumulated less nitrogen than did unsupported plants, suggesting a less efficient uptake of this nutrient. The ability to plastically modify growth and allocation patterns in response to trellis found in some lianas may constitute an important aspect of their ecological strategy. In other species, however, the switch to climbing may incur physiological costs resulting in lower growth.

BACKGROUND AND AIMS:The reliance on external support by lianas has been hypothesized to imply a reduction in the biomass cost of stem construction and root anchorage, and an increased investment in leaves, relative to self-supporting plants. These evolutionary trade-offs have not been adequately tested in an ontogenetic context and on the whole-plant scale. Moreover, the hypothesis may be extended to other potentially limiting resources, such as nitrogen (N.). METHODS:Plants belonging to five con-familiar pairs of temperate liana/shrub species were cultivated in 120 L barrels and sequentially harvested over up to three growing seasons. To account for the ontogenetic drift, organ biomass and nitrogen fractions were adjusted for plant biomass and N pool, respectively. KEY RESULTS:Lianas invested, on average, relatively less biomass in the root fraction in comparison with shrubs. This was offset by only insignificant increases in leaf or stem investment. Even though liana stems and roots showed higher N concentration in comparison with shrubs, plant N distribution was mostly driven by, and largely matched, the pattern of biomass distribution. Lianas also showed a greater relative growth rate than shrubs. The differences between the growth forms became apparent only when ontogenetic drift was controlled for. These results were confirmed regardless of whether reproductive biomass was included in the analysis. CONCLUSIONS:Our results suggest that temperate lianas, in spite of their diverse, species-specific resource distribution patterns, preferentially allocate resources to above-ground organs at the expense of roots. By identifying this trade-off and demonstrating the lack of a general trend for reduction in stem investment in lianas, we significantly modify the prevailing view of liana allocation strategies and evolutionary advantages. Such a resource distribution pattern, along with the cheap unit leaf area and stem unit length construction, situates lianas as a group close to the fast acquisition/rapid growth end of the life strategy spectrum.

Aims While patterns of variation in woody plant aboveground traits related to disturbance and resource availability in savanna ecosystems are fairly well understood, dimensions of variation in belowground traits remain poorly understood. We investigate how sapling coarse root (>2 mm diameter) traits and belowground storage organs vary with respect to differing disturbance regimes (i.e. fire or herbivore dominated) and soil fertility in Southern African savannas. Methods We examined how sapling rooting strategies (type of rooting system, storage organs, root allocation) as well as root total non-structural carbohydrate concentrations [TNC], root N and C:N of 69 common woody savanna and forest species at 16 sites vary with respect to differing disturbance regimes and soil fertility. Results We found that root [TNC] and the occurrence of storage organs were highest while root N was lowest for species growing at fire-driven sites, compared to herbivore-driven sites and competition-controlled forest species. Allocation to belowground biomass was higher at fire-dominated sites but root volume:stem basal area did not differ between disturbance regimes. None of the measured traits were found to differ between nutrient rich clayey soils and nutrient poor sandy soils. Conclusions Our results suggest that disturbance related controls are important drivers of savanna belowground traits.