Temperate Tree Research Articles

Patterns and controlling factors of decomposition in distal shoot systems by branch order across 10 temperate tree species

Patterns and controlling factors of decomposition in distal shoot systems by branch order across 10 temperate tree species

Rapid estimates of leaf litter chemistry using reflectance spectroscopy

Measuring the chemical traits of leaf litter is important for understanding plants’ influence on nutrient cycles, including through nutrient resorption and litter decomposition, but conventional leaf trait measurements are often destructive and labor-intensive. Here, we develop and evaluate the performance of partial least-squares regression models that use reflectance spectra of intact or ground leaves to estimate leaf litter traits, including carbon and nitrogen concentration, carbon fractions, and leaf mass per area (LMA). Our analyses included more than 300 samples of senesced foliage from 11 species of temperate trees, including both needleleaf and broadleaf species. Across all samples, we could predict each trait with moderate-to-high accuracy from both intact-leaf litter spectra (validation R2 = 0.543–0.941; %root mean squared error (RMSE) = 7.49–18.5) and ground-leaf litter spectra (validation R2 = 0.491–0.946; %RMSE = 7.00–19.5). Notably, intact-leaf spectra yielded better predictions of LMA. Our results support the feasibility of building models to estimate multiple chemical traits from leaf litter of a range of species. In particular, intact-leaf spectral models allow non-destructive trait estimation in a matter of seconds, which could enable researchers to measure the same leaves over time in studies of nutrient resorption.

Perspectives on the adaptation of Japanese plum-type cultivars to reduced winter chilling in two regions of Spain.

Japanese plum, like other temperate fruit tree species, has cultivar-specific temperature requirements during dormancy for proper flowering. Knowing the temperature requirements of this species is of increasing interest due to the great genetic variability that exists among the available Japanese plum-type cultivars, since most of them are interspecific hybrids. The reduction of winter chilling caused by climate change is threatening their cultivation in many regions. In this work, the adaptation perspectives of 21 Japanese plum-type cultivars were analyzed in two of the main plum-growing regions in Spain, Badajoz and Zaragoza, to future climate conditions. Endodormancy release for subsequent estimation of chilling and heat requirements was determined through empirical experiments conducted during dormancy at least over two years. Chill requirements were calculated using three models [chilling hours (CH), chilling units (CU) and chilling portions (CP)] and heat requirements using growing degree hours (GDH). Chilling requirements ranged 277-851 CH, 412-1,030 CU and 26-51 CP, and heat requirements ranged from 4,343 to 9,525 GDH. The potential adaption of the cultivars to future warmer conditions in both regions was assessed using climate projections under two Representative Concentration Pathways (RCP), RCP4.5 (effective reduction of greenhouse gas emissions) and RCP8.5 (continuous increase in greenhouse gas emissions), in two time horizons, from the middle to the end of 21st century, with temperature projections from 15 Global Climate Models. The probability of satisfying the estimated cultivar-specific chilling requirements in Badajoz was lower than in Zaragoza, because of the lower chill availability predicted. In this region, the cultivars analyzed herein may have limited cultivation because the predicted reduction in winter chill may result in the chilling requirements not being successfully fulfilled.

Phenomics and transcriptomic profiling of fruit development in distinct apple varieties

Apple is one of the most economically important and popular temperate fruit trees. The domestication of apple has resulted in substantial phenotypic differences, particularly between wild and cultivated varieties. However, the relationship between gene expression and phenotypic variations in apple remains poorly understood. Here, we present a comprehensive dataset featuring five distinct apple varieties, including two wild varieties and three representative cultivated varieties. The dataset comprises of both phenomics data, encompassing twelve fruit quality-related traits continuously measured over two years, and transcriptomic data obtained at different developmental stages with three biological replicates. We performed basic quality control process, gene expression normalization and differential gene expression analysis to demonstrate the utility and reliability of the dataset. Our findings indicate that gene expression strongly related with phenotypic variations in apple. This dataset serves as a valuable resource, encompassing phenomics and transcriptomic data in multiple formats, thereby facilitating further exploration of the relationships between gene expression and phenotypic traits in apple.

Coordinated hydraulic traits influence the two phases of time to hydraulic failure in five temperate tree species differing in stomatal stringency.

Worldwide, forests are increasingly exposed to extreme droughts causing tree mortality. Because of the complex nature of the mechanisms involved, various traits have been linked to tree drought responses with contrasting results. This may be due to species-specific strategies in regulating water potential, a process that unfolds in two distinct phases: a first phase until stomatal closure, and a second phase until reaching lethal xylem hydraulic thresholds. We conducted dry-down experiments with five broadleaved temperate tree species differing in their degree of isohydry to estimate the time to stomatal closure (tsc) and subsequent time to critical hydraulic failure (tcrit). We measured various traits linked to tree drought responses, such as the water potentials at turgor loss point (Ptlp), stomatal closure (Pgs90), and 12%, 50% and 88% loss of xylem hydraulic conductance (P12, P50, P88), hydraulic capacitance (C), minimum leaf conductance (gmin), hydroscape area (HSA) and hydraulic safety margins (HSM). We found that Pgs90 followed previously recorded patterns of isohydry and was associated with HSA. Species ranked from more to less isohydric in the sequence Acer pseudoplatanus<Betula pendula<Tilia cordata<Sorbus aucuparia<Fagus sylvatica. Their degree of isohydry was associated with leaf safety (Ptlp and gmin), drought avoidance (C) and tsc, but decoupled from xylem safety (HSM and P88) and tcrit. Regardless of their stomatal stringency, species with wider HSM and lower P88 reached critical hydraulic failure later. We conclude that the duration of the first phase is determined by stomatal regulation, while the duration of the second phase is associated with xylem safety. Isohydry is thus linked to water use rather than to drought survival strategies, confirming the proposed use of HSA as a complement to HSM for describing plant drought responses before and after stomatal closure.

Spatial phylogenetics of Fagales: Investigating drivers of temperate forest distributions

AbstractAimQuantifying the phylogenetic diversity of temperate trees is essential for understanding the processes that have shaped the modern distribution of temperate broadleaf forest and other major forest biomes. Here, we focus on Fagales, an iconic member of forests worldwide, to uncover global diversity and endemism patterns and investigate the distribution of root nodule symbiosis (RNS), an important morphological specialisation in this clade, as a key factor behind these patterns.LocationGlobal.TaxonFagales.MethodsWe combined phylogenetic data covering 60.2% of living species, fine‐scale distribution models covering 90% of species, and nodulation data covering all species to investigate the distribution of species richness and phylogenetic diversity at fine spatial scales compared to the distribution of RNS. We identify abiotic environmental factors associated with RNS and with Fagales diversity in general.ResultsWe find the highest species richness in temperate east Asia, eastern North America, and equatorial montane regions of Asia and Central America. By contrast, relative phylogenetic diversity (RPD) is highest at higher latitudes, where RNS also predominates. We found a strong spatial structuring of regionalisations of Fagales floras, reflecting distinct Northern and Southern Hemisphere floras (except a unique Afro‐Boreal region), each with distinct RNS‐environment relationships.Main ConclusionsAlthough species richness and phylogenetic regionalisation for Fagales accord well with traditional biogeographic concepts for temperate forests, this is not the case for RPD. RNS is almost universal in the highest RPD regions, which may reflect ecological filtering promoting RNS in these regions. Our results highlight the utility of global‐scale, clade‐specific spatial phylogenetics and its utility for understanding drivers of diversity in species‐rich clades.

Species-specific responses of C and N allocation to N addition: evidence from dual 13C and 15N labeling in three tree species

Soil nitrogen (N) availability affects plant carbon (C) utilization. However, it is unclear how various tree functional types respond to N addition in terms of C assimilation, allocation, and storage. Here, a microcosm experiment with dual 13C and 15N labeling was conducted to study the effects of N addition (i.e., control, 0 g N kg−1; moderate N addition, 1.68 g N kg−1; and high N addition, 3.36 g N kg−1 soil) on morphological traits, on changes in nonstructural carbohydrates (NSC) in different organs, as well as on C and N uptake and allocation in three European temperate forest tree species (i.e., Acer pseudoplatanus, Picea abies and Abies alba). Our results demonstrated that root N uptake rates of the three tree species increased by N addition. In A. pseudoplatanus, N uptake by roots, N allocation to aboveground organs, and aboveground biomass allocation significantly improved by moderate and high N addition. In A. alba, only the high N addition treatment considerably raised aboveground N and C allocation. In contrast, biomass as well as C and N allocation between above and belowground tissues were not altered by N addition in P. abies. Meanwhile, NSC content as well as C and N coupling (represented by the ratio of relative 13C and 15N allocation rates in organs) were affected by N addition in A. pseudoplantanus and P. abies but not in A. alba. Overall, A. pseudoplatanus displayed the highest sensitivity to N addition and the highest N requirement among the three species, while P. abies had a lower N demand than A. alba. Our findings highlight that the responses of C and N allocation to soil N availability are species-specific and vary with the amount of N addition.

Drought Responses in Three Apple Cultivars Using an Autonomous Sensor-based Irrigation System

Irrigation decision support systems evolving in the domestic temperate tree fruit production industry incorporate measures of soil moisture status, which diverges from classic physiological indicators of edaphic stress. This study used an autonomous sensor-based irrigation system to impose a water deficit (soil matric potential targets of –25, –40, –60, and –80 kPa) on ‘Autumn Gala’, ‘CrimsonCrisp’, and ‘Golden Delicious’ apple (Malus domestica) cultivars grafted to ‘Budagovsky 9’ rootstock in the greenhouse (n = 60). It was hypothesized that relationships between physiological plant function, assessed via infrared gas exchange and chlorophyll fluorescence, and the soil matric potential may be used to advance emerging irrigation decision support systems. Complications arising from defoliation by day 11 at –60 and –80 kPa indicate the generation of substrate-specific soil–water relationships in research applications of autonomous sensor-based irrigation systems. ‘Autumn Gala’ carbon assimilation rates at –80 kPa declined from day 0 to day 8 (9.93 and 5.86 μmol⋅m–2⋅s–1 carbon dioxide), whereas the transpiration rate was maintained, potentially reducing observed defoliation as other cultivars increased transpiration to maintain carbon assimilation. Correlation matrices revealed Pearson’s r ≤ |0.43| for all physiological metrics considered with soil matric potential. Nevertheless, exploratory regression analysis on predawn leaf water potential, carbon assimilation, transpiration, stomatal conductance, and nonphotochemical quenching exposed speculatively useful data and data shapes that warrant additional study. Nonlinear piecewise regression suggested soil matric potential may useful as a predictor for the rate of change in predawn leaf water potential upon exposure to a water deficit. The critical point bridging the linear spans, –30.6 kPa, could be useful for incorporating in emerging irrigation decision support systems.

Foliar spectra accurately distinguish most temperate tree species and show strong phylogenetic signal.

Spectroscopy is a powerful remote sensing tool for monitoring plant biodiversity over broad geographic areas. Increasing evidence suggests that foliar spectral reflectance can be used to identify trees at the species level. However, most studies have focused on only a limited number of species at a time, and few studies have explored the underlying phylogenetic structure of leaf spectra. Accurate species identifications are important for reliable estimations of biodiversity from spectral data. Using over 3500 leaf-level spectral measurements, we evaluated whether foliar reflectance spectra (400-2400 nm) can accurately differentiate most tree species from a regional species pool in eastern North America. We explored relationships between spectral, phylogenetic, and leaf functional trait variation as well as their influence on species classification using a hurdle regression model. Spectral reflectance accurately differentiated tree species (κ = 0.736, ±0.005). Foliar spectra showed strong phylogenetic signal, and classification errors from foliar spectra, although present at higher taxonomic levels, were found predominantly between closely related species, often of the same genus. In addition, we find functional and phylogenetic distance broadly control the occurrence and frequency of spectral classification mistakes among species. Our results further support the link between leaf spectral diversity, taxonomic hierarchy, and phylogenetic and functional diversity, and highlight the potential of spectroscopy to remotely sense plant biodiversity and vegetation response to global change.

The role of dispersal limitation in the forest biome shifts of Europe in the last 18,000 years

AbstractAimHow the ability of plants to move towards newly favourable habitats (dispersal limitation) impacts the change of biome distribution and transition under fast climate warming is still debated. Analysing vegetation change in the past may help to clarify the relative importance of underlying ecological processes such as climate, biotic interactions, and dispersal. In this study, we investigated how dispersal limitation affected the distribution of European forests in the last 18,000 years.LocationSouthern and Central Europe.TaxonSpermatophyta.MethodsUsing the LPJ‐GM 2.0 model (an extension of LPJ‐GUESS), we simulated European vegetation from the end of the Last Glacial Maximum (18.5 ka) to the current time (0 ka). Using biome reconstructions from pollen data as reference, we compared the performance of two dispersal modes: with no migration constraints or seed limitation (free dispersal mode), and with plant establishment depending on seed dynamics and dispersal (dispersal limitation mode).ResultsThe model run, including migration processes, was better at capturing the post‐glacial expansion of European temperate forests (and the longer persistence of boreal forests) than the setting assuming free dispersal, especially during periods of rapid warming. This suggests that a number of (temperate) tree taxa experienced delayed occupancy of climatically suitable habitats due to a limited dispersal capacity, i.e., post‐glacial migration lags.Main ConclusionsOur results show that including migration processes in model simulations allows for more realistic reconstructions of forest patterns under rapid climate change, with consequences for future projections of carbon sequestration and climate reconstructions with vegetation feedback, assisted migration and forest conservation.

Stem decomposition of temperate tree species is determined by stem traits and fungal community composition during early stem decay

Abstract Dead trees are vital structural elements in forests playing key roles in the carbon and nutrient cycle. Stem traits and fungal community composition are both important drivers of stem decay, and thereby affect ecosystem functioning, but their relative importance for stem decomposition over time remains unclear. To address this issue, we used a common garden decomposition experiment in a Dutch larch forest hosting fresh logs from 13 common temperate tree species. In total, 25 fresh wood and bark traits were measured as indicators of wood accessibility for decomposers, nutritional quality and chemical or physical defence mechanisms. After 1 and 4 years of decay, we assessed the richness and composition of wood‐inhabiting fungi using amplicon sequencing and determined the proportional wood density loss. Average proportional wood density loss for the first year was 18.5%, with further decomposition occurring at a rate of 4.3% year−1 for the subsequent 3 years across tree species. Proportional wood density loss varied widely across tree species in the first year (8.7–24.8% year−1) and subsequent years (0–11.3% year−1). The variation was directly driven by initial wood traits during the first decay year, then later directly driven by bark traits and fungal community composition. Moreover, bark traits affected the composition of wood‐inhabiting fungi and thereby indirectly affected decomposition rates. Specifically, traits promoting resource acquisition of the living tree, such as wide conduits that increase accessibility and high nutrient concentration, increased initial wood decomposition rates. Fungal community composition, but not fungal richness explained differences in wood decomposition after 4 years of exposure in the field, where fungal communities dominated by brown‐rot and white‐rot Basidiomycetes were linked to higher wood decomposition rate. Synthesis: Understanding what drives deadwood decomposition through time is important to understand the dynamics of carbon stocks. Here, using a tailor‐made experimental design in a temperate forest setting, we have shown that stem trait variation is key to understanding the roles of these drivers; initially, wood traits explained decomposition rates while subsequently, bark traits and fungal decomposer composition drove decomposition rates. These findings inform forest management with a view to selecting tree species to promote carbon storage.

Genome-wide identification analysis of the 4-Coumarate: CoA ligase (4CL) gene family expression profiles in Juglans regia and its wild relatives J. Mandshurica resistance and salt stress

Persian walnut (Juglans regia) and Manchurian walnut (Juglans mandshurica) belong to Juglandaceae, which are vulnerable, temperate deciduous perennial trees with high economical, ecological, and industrial values. 4-Coumarate: CoA ligase (4CL) plays an essential function in plant development, growth, and stress. Walnut production is challenged by diverse stresses, such as salinity, drought, and diseases. However, the characteristics and expression levels of 4CL gene family in Juglans species resistance and under salt stress are unknown. Here, we identified 36 Jr4CL genes and 31 Jm4CL genes, respectively. Based on phylogenetic relationship analysis, all 4CL genes were divided into three branches. WGD was the major duplication mode for 4CLs in two Juglans species. The phylogenic and collinearity analyses showed that the 4CLs were relatively conserved during evolution, but the gene structures varied widely. 4CLs promoter region contained multiply cis-acting elements related to phytohormones and stress responses. We found that Jr4CLs may be participated in the regulation of resistance to anthracnose. The expression level and some physiological of 4CLs were changed significantly after salt treatment. According to qRT-PCR results, positive regulation was found to be the main mode of regulation of 4CL genes after salt stress. Overall, J. mandshurica outperformed J. regia. Therefore, J. mandshurica can be used as a walnut rootstock to improve salt tolerance. Our results provide new understanding the potential functions of 4CL genes in stress tolerance, offer the theoretical genetic basis of walnut varieties adapted to salt stress, and provide an important reference for breeding cultivated walnuts for stress tolerance.

Cambium Reactivation Is Closely Related to the Cell-Cycle Gene Configuration in Larix kaempferi.

Dormancy release and reactivation in temperate trees are mainly controlled by temperature and are affected by age, but the underlying molecular mechanisms are still unclear. In this study, we explored the effects of low temperatures in winter and warm temperatures in spring on dormancy release and reactivation in Larix kaempferi. Further, we established the relationships between cell-cycle genes and cambium cell division. The results showed that chilling accelerated L. kaempferi bud break overall, and the longer the duration of chilling is, the shorter the bud break time is. After dormancy release, warm temperatures induced cell-cycle gene expression; when the configuration value of the cell-cycle genes reached 4.97, the cambium cells divided and L. kaempferi reactivated. This study helps to predict the impact of climate change on wood production and provides technical support for seedling cultivation in greenhouses.

Temperate woody species across the angiosperm phylogeny acquire tolerance to water deficit stress during the growing season.

Understanding the capacity of temperate trees to acclimate to limited soil water has become essential in the face of increasing drought risk due to climate change. We documented seasonal - or phenological - patterns in acclimation to water deficit stress in stems and leaves of tree species spanning the angiosperm phylogeny. Over 3 yr of field observations carried out in two US arboreta, we measured stem vulnerability to embolism (36 individuals of 7 Species) and turgor loss point (119 individuals of 27 species) over the growing season. We also conducted a growth chamber experiment on 20 individuals of one species to assess the mechanistic relationship between soil water restriction and acclimation. In three-quarters of species measured, plants became less vulnerable to embolism and/or loss of turgor over the growing season. We were able to stimulate this acclimatory effect by withholding water in the growth chamber experiment. Temperate angiosperms are capable of acclimation to soil water deficit stress, showing maximum vulnerability to soil water deficits following budbreak and becoming more resilient to damage over the course of the growing season or in response to simulated drought. The species-specific tempo and extent of this acclimatory potential constitutes preadaptive climate change resilience.

Discrepancies in methodologies to determine chill requirements in temperate fruit trees constrain guidelines for future plantings in a global warming context

Temperate fruit trees enter a dormant state and resume growth development once cultivar-specific chill requirements (CR) are fulfilled. Sustainable fruit production requires crucial knowledge of the CR of the varieties to predict the adaptability of crops to future climate conditions. Experimental methodologies based on exposing shoots to forcing conditions, and statistical methods based on analysing long-term phenology records, such as Partial Least Square regression (PLS), are commonly used to estimate CR. To demonstrate the importance of unifying methodologies, we compared CR estimations from six approaches that included bud weight increase (30 % and 50 %), different percentages of bud break (30 %, 50 % and 90 %) after forcing conditions, and PLS regression. The CR of four almond and four apricot cultivars were experimentally and statistically determined in Murcia, South-Eastern Spain. Comparisons between approaches revealed that the choice of the method had a significant effect on the estimates. Large variation in CR estimates was found between PLS regression and the experimental methodologies in some late-flowering cultivars. To assess the adaptation potential of a certain cultivar to future climate conditions based on the CR estimates obtained through the different methodologies, we compared the probability of fulfilling cultivar-specific CR's under two global warming scenarios. Projections revealed a marked decreasing trend of chill availability in the region, which is already threatening the cultivation of genotypes with high CR. This study demonstrates that the variability of methodologies used to quantify CR has an important impact on assessing the probability of a successful adaptation. CR obtained by a certain methodology suggest that insufficient chill might be an increasingly common situation for some cultivars, potentially resulting in production risks. Conversely, other methodologies appear to indicate a more favourable performance for the same cultivar. Implementing standardized methods and metrics is essential to ensure more reliable information regarding the prospects for future crop adaptation.

Climate Change Challenges in Temperate and Sub-Tropical Fruit Tree Cultivation

In the last few years, the world has experienced the impacts of climate change, such as elevated mean annual temperature, extreme weather events, drought, etc. Among living organisms, perennial plant species are the ones mostly exposed to climate change impacts, as they may experience different extreme events within the same year, such as flooding during some periods and drought in summer months, extremely low temperatures in winter but excessively high temperatures in summer, etc. Climate change affects a range of physiological functions of temperate fruit and nut tree species, such as their phenophases, bud dormancy release and vernalization, pollination and fruit set, fruit growth and quality, as well as bud sprouting and growth initiation. Besides these, the impact of climate change on pests, diseases, and weeds may generate significant negative interactions with tree physiology, threatening food production, food safety, and human welfare. In the present manuscript, a general aspect of climate change impacts on fruits’ and nut trees’ physiological functions is described and commented on.

The clockwork of spring: bud dormancy timing as a driver of spring leaf-out in temperate deciduous trees

The initiation of spring leaf-out is a critical determinant of the growing season in trees, affecting primary production and species interactions in forest ecosystems. Variations in the timing of leaf-out among tree species are linked to their differential progression of bud dormancy. However, identifying reliable markers for bud dormancy has been challenging, leaving the connection between the timing of autumn leaf senescence, bud dormancy and spring leaf-out unclear. To test whether species initiating dormancy release earlier also exhibit earlier leaf senescence in autumn and earlier leaf-out in spring, we estimated the dates of peak dormancy depth (PDD), senescence timing and spring leaf-out across various species, locations and experimental conditions in Central Europe. PDD was defined as the date when the maximum thermal sum was required for leaf-out, whereas leaf senescence was assessed through the decrease in leaf greenness. Our findings reveal that PDD timing is a more accurate predictor of species-level differences in spring leaf-out dates than the timing of leaf senescence, the latter being a poor proxy for PDD. The observed temporal asynchrony between PDD and senescence was linked to dormancy induction showing species-specific sensitivity to temperature variations. Conversely, the timing of leaf senescence showed a consistent reaction to temperature changes across all species. These findings suggest that the physiological processes within buds and leaves during autumn are governed by distinct environmental cues, with the bud dormancy process serving as a more reliable predictor of spring phenological differences among forest tree species than does autumn leaf senescence.

The sensitivity of root water uptake to cold root temperature follows species-specific upper elevational distribution limits of temperate tree species.

Physiological water stress induced by low root temperatures might contribute to species-specific climatic limits of tree distribution. We investigated the low temperature sensitivity of root water uptake and transport in seedlings of 16 European tree species which reach their natural upper elevation distribution limits at different distances to the alpine treeline. We used 2H-H2O pulse-labelling to quantify the water uptake and transport velocity from roots to leaves in seedlings exposed to constant 15°C, 7°C or 2°C root temperature, but identical aboveground temperatures between 20°C and 25°C. In all species, low root temperatures reduced the water transport rate, accompanied by reduced stem water potentials and stomatal conductance. At 7°C root temperature, the relative water uptake rates among species correlated positively with the species-specific upper elevation limits, indicating an increasingly higher sensitivity to lower root zone temperatures, the lower a species'natural elevational distribution limit. Conversely, 2°C root temperature severely inhibited water uptake in all species, irrespective of the species'thermal elevational limits. We conclude that low temperature-induced hydraulic constraints contribute to the cold distribution limits of temperate tree species and are a potential physiological cause behind the low temperature limits of plant growth in general.

Drought effects on growth and density of temperate tree regeneration under different levels of nitrogen deposition

Temperate forests in Central Europe suffer from climate change-induced productivity and vitality reductions and increased tree mortality. Most field work assessing climate change effects in forests refers to mature trees and does not cover interaction between climate change and nitrogen deposition. Here we show in a study of 54 forest sites representing different combinations of climatic conditions and atmospheric nitrogen deposition across Germany that nitrogen significantly affects the drought tolerance of tree regeneration in the field. We compared shoot length increment and regeneration density of Central Europe’s naturally most dominant tree species, European beech (Fagus sylvatica) with three species, which are discussed as potentially more drought-tolerant replacement tree species for climate change adaptation of forestry. Growth of beech was reduced with increasing nitrogen deposition identifying high reactive nitrogen loads as an additional threat for this species, but between-site variation of beech growth was not dependent on climatic parameters despite growth reductions after the drought summer 2018 across all study sites. Remarkably, the only species in the between-site comparisons where low growth was attributable to dry climate was Douglas fir (Pseudotsuga menziesii), as shoot length increment was reduced at sites with dry spring climate. Silver fir (Abies alba) showed increasing growth with increasing nitrogen deposition in combination with increasing drought, probably due to reduced ammonium uptake. Shoot length increment in sessile oak (Quercus petraea) was not affected by the between-site variation of either drought or nitrogen. Our results (combined with results of regeneration density) indicate that high atmospheric nitrogen deposition, which is primarily found in regions with intensive livestock farming in Central Europe, modifies the climate change response of temperate tree species. Interaction of drought and nitrogen deposition should thus be addressed in the debate on climate change adaptation of forestry.

Climate warming advances phenological sequences of Aesculus hippocastanum

Warming-induced shifts in plant phenology have been frequently reported in recent decades. Previous studies primarily focused on isolated phenological events; however, the entire phenological sequences of a given species were rarely examined simultaneously, limiting our knowledge about aggregate life history response to climate warming. Here, using 210,000 in situ phenological observations across 2493 European sites during 1980–2015, we examined the responses of four typical phenological events (leaf unfolding, flowering, fruiting, and leaf coloring) of Aesculus hippocastanum (horse chestnut), a common temperate tree species, to climate warming. We found that all four phenophases of A. hippocastanum advanced, i.e., the entire life cycle during the year shifted earlier with climate warming. However, the early-season phenophases (leaf unfolding and flowering) advanced faster than late-season phenophases (leaf coloring and fruiting), leading to the extension of both growing season (from leaf unfolding to leaf coloring) and reproductive season (from flowering to fruiting). The extension rate of the reproductive season was slightly greater than that of the growing season. This implied that under warming climate, more time was devoted to both vegetative growth and reproduction, particularly the latter one, in A. hippocastanum. In addition, significantly positive correlations were observed among four phenological events, particularly for pairwise consecutive events, indicating that the responses of phenological sequences to warming were highly correlated. By simultaneously studying the responses of four key phenological events to climate warming, our findings demonstrate the differences in the phenological plasticity among phenological sequences in A. hippocastanum, deepening our understanding about the impact of climate warming on life history rather than examining individual phenological events in isolation.