Tree Physiology

Compound droughts, i.e. the co-occurrences of heat and drought, represent a serious challenge for temperate forest trees leading to significant losses in forest biomass. We studied the physiological response of Norway spruce (Picea abies) saplings to heat and drought individually, and in combination. Continuous measurements of leaf gas exchange and VOC emission allowed us to identify fast-response reactions, while discrete VOC and root exudate samplings added qualitative information on compositional changes. Additionally, we used 13CO2 and 2H2O label pulses to investigate C-allocation and root water uptake in response to stress. Heat as well as drought reduced assimilation rates in the saplings, whereas transpiration, leaf VOC emission and root exudation rates increased in response to heat. Drought alone increased VOC emission but decreased exudation rates. Combined heat and drought triggered an amplified response in both processes despite negative net CO2 assimilation rates. Label incorporation showed compromised water uptake capacity of drought-stressed plants and illustrated de novo C-allocation to VOC emission and root exudates. The results point at the high susceptibility of Norway spruce saplings to drought and heat. Combined stress resulted in synergistic responses in VOC emissions and root exudates, showing the detrimental effect of compound droughts on Norway spruce. HighlightIn this study, we found synergistic effects of heat and drought on carbon losses from leaf VOC emission and root exudates despite negative assimilation rates in Norway spruce saplings.

The article introduces a new Forest Stress Index (ISF) based on a plant hydraulic modelling approach rather than classical climatic drought indices. Unlike other index like scPDSI or SPEI, ISF is grounded in xylem embolism dynamics simulated with the mechanistic SurEau model. The goal is to better link climatic anomalies to tree physiological functioning and mortality risk. ISF is defined using a locally adapted ideotype characterized by an optimal P50 value under a reference hydraulic functioning threshold. Simulations are performed across Europe and France using multiple climate datasets. The index is robust to model parameterization choices and assumptions about plant functional traits. Results show strong spatial and temporal consistency and significant correlations with SPEI and scPDSI. However, ISF more strongly highlights extreme drought years and exhibits a more skewed distribution. Future projections under SSP5-8.5 indicate a widespread increase in hydraulic stress with strong regional contrasts. Overall, ISF provides a mechanistic and complementary drought indicator more directly linked to forest mortality processes.

Xylem and bark properties influence tree growth and drought resistance, yet their functional coordination and their environmental drivers remain unclear. We assessed xylem-bark coordination in branches of eight temperate woody Rosaceae species spanning different ecological preferences. We quantified xylem hydraulic efficiency and safety alongside bark traits governing permeability, hygroscopic water exchange, water storage, and anatomy, and evaluated phylogenetic signal and climatic associations. Bark water vapor conductance (Gbark) increased with maximum xylem hydraulic conductivity (Kh) and with xylem water potential at 50% loss of conductivity (P50), indicating species with more efficient but more embolism-vulnerable xylem developed more permeable bark. Species with higher Gbark showed reduced hygroscopic absorption time, consistent with faster rehydration from atmospheric water vapor. Both Gbark and P50 were phylogenetically conserved and covaried with climatic factors, namely air temperature, vapor pressure deficit (VPD), and isothermality. Species from warmer, high-VPD climates with greater diurnal temperature variability combined higher bark permeability with more vulnerable xylem, implying a shift from embolism avoidance to embolism tolerance strategies. Overall, xylem and bark hydraulics in Rosaceae evolved in concert along diurnal and annual gradients of evaporative demand, showing that drought resistance in woody angiosperms cannot be understood without considering bark traits alongside xylem function. Graphical abstract O_FIG O_LINKSMALLFIG WIDTH=200 HEIGHT=104 SRC="FIGDIR/small/730605v1_ufig1.gif" ALT="Figure 1"> View larger version (41K): org.highwire.dtl.DTLVardef@14872c2org.highwire.dtl.DTLVardef@1662c8corg.highwire.dtl.DTLVardef@f6b4aaorg.highwire.dtl.DTLVardef@cf1aef_HPS_FORMAT_FIGEXP M_FIG C_FIG Caption: This study shows that xylem and bark in Rosaceae species form an integrated functional system in which xylem hydraulic safety, efficiency, and bark permeability are jointly tuned along diurnal and annual gradients of air temperature and evaporative demand.

Trees are routinely pruned to mitigate the risk of wind damage, but there are few studies examining changes in wind loads after pruning, especially for large conifers. In this study, ten Colorado spruces (Picea pungens) were monitored before and after a series of pruning treatments. Trees were pruned to raise or thin crowns over a range of severities between 0% and 40%. Wind-induced bending moments were measured using two calibrated displacement probes installed orthogonally on the lower stem of each tree. Using a hierarchical Bayesian model, the relationship between maximum wind speeds and bending moments was quantified, consistent with theoretical and empirical expectations, as a non-linear power law. Random intercepts for model coefficients were used to account for individual variability in aerodynamic behavior among experimental trees, and predictions were made using the median response marginalized over the observed trees. The modeled relationship between wind speeds and bending moments was physically reasonable and like existing measurements with scaling exponents below two. Despite considerable variation among experimental trees, the aerodynamic behavior of trees, as indicated by model coefficients, was not clearly altered by pruning treatments, and, correspondingly, model predictions of bending moments over the range of observed wind speeds remained similar for all pruning treatments. Ultimately, the study yielded weak evidence for a change in bending moments following conventional pruning treatments for Colorado spruce, and the practical value of pruning to mitigate risk appeared limited for the studied conditions. Highlights- Wind loads were monitored on large Colorado spruce after crown raising and thinning - A hierarchical Bayesian model quantified wind speed and bending moment power laws - Negligible change in bending moments was found for all pruning types and severities - Conventional pruning methods may not mitigate risk for Colorado spruce

The sequence of leaf physiological thresholds underlying plant responses to water deficit is thought to be functionally coordinated; yet, to what extent this coordination is maintained across genotypes and environments remains poorly documented at the intraspecific level. We characterized the sequence of stomatal closure, turgor loss and xylem embolism in the leaves of two genotypes of the riparian species Populus nigra (DRA-038 vs. PG-31) subjected to control, additional nitrogen or additional potassium treatments. Under control conditions, embolism measurements using the optical vulnerability method showed that DRA-038 was more vulnerable than PG-31, in agreement with measurements performed on stems with the reference Cavitron method. Stomatal closure consistently preceded xylem embolism, while bulk leaf turgor loss was typically observed once xylem embolism had already reached 50%. Hydraulic thresholds responded to treatments in a genotype-dependent manner, the intrinsically more vulnerable genotype DRA-038 being typically more plastic. However, despite variations across genotypes and treatments, the trait sequence remained tightly coordinated such that stomatal safety margins (SSMs) remained virtually null. These findings support a strong mechanistic integration of leaf hydraulic thresholds in poplar across genetic units and varying environments, questioning whether to favour intrinsic tolerance or plastic capacities in breeding future drought-tolerant genotypes.

Bacterial canker remains a major constraint affecting apricot production in South-East of France. It is primarily caused by Pseudomonas syringae, a Gram-negative bacterium, many strains of which exhibit ice nucleation activity. By promoting ice formation at relatively high subzero temperatures, ice nucleation-active bacteria may facilitate tissue disruption and pathogen entry. Concurrently, climate-driven shifts toward warmer winter-spring periods have advanced flowering phenology, increasing exposure to late frost events. Despite breeders having developed less susceptible varieties to bacterial canker and early flowering varieties, the link between these traits and frost sensitivity, an emerging risk in this location, remains unresolved. Here, we have evaluated the links between canker susceptibility and frost sensitivity using three cultivar pairs contrasting in disease response and flowering time. Ice nucleation temperature was measured in excised buds under controlled conditions throughout the frost-risk period, alongside field-based diameter variation monitoring over two years. Disease susceptibility (P < 0.001), phenology (P = 0.003), varieties (P < 0.001), locations (P < 0.001), and sampling date (P < 0.001) significantly affected nucleation temperature, whereas epiphytic bacterial abundance and xylem vessel diameter did not. Trees froze at higher temperatures in situ than in laboratory assays (1 to -2{degrees}C versus -3 to 4{degrees}C, respectively), indicating strong environmental modulation of freezing processes beyond Psy-like bacterial activity, which is reflected in contrasting disease susceptibilities (P < 0.001) and precocities (P < 0.001). These results shed light on the complexity of the freezing process in trees under natural conditions. We discuss the potential roles of microclimatic conditions and alternative ice nucleation sources beyond Psy-like bacteria in driving these physiological processes.

We tested whether seasonal plasticity in photosystem II (PSII) heat tolerance aligns with leaf phenology in tropical trees, comparing evergreen and deciduous species across the wettopostwet transition in a seasonally dry tropical forest of the central Western Ghats, India. This transition, preceding droughtinduced senescence, represents the key window to assess true unstressed thermal plasticity. PSII thermal traits: damage onset (T5), damage midpoint (T50), and temperature between damage onset to full loss (T95-T5), decline width (DW) were quantified in 27 cooccurring species during the wet (27.5{square}{degrees}C) and postwet (31.6{square}{degrees}C) periods. Contrary to phenologybased predictions, PSII plasticity was not structured by leaf habit or successional status. Both T5 (+1.7{square}{degrees}C) and T50 (+0.9{square}{degrees}C) increased significantly across seasons, but responses were speciesspecific, with evergreen and deciduous trees acclimating similarly. The preventionversusforbearance tradeoff (T5 - DW relationship) remained conserved, though leaf habits diverged under postwet conditions. Thermal safety margins based on T50 were large, but T5 revealed vulnerable latesuccessional evergreens (Saraca asoca, Ficus spp.) and Careya arborea. These results show that PSII thermotolerance regulation operates largely independently of droughtavoidance phenology, indicating species identity and not leaf habit drives plasticity in PSII thermal response in seasonally dry tropical forests.

Due to climate change, extreme weather events such as droughts are becoming more frequent and intense. This has a profound impact on plant performance and ecological interactions, including those involving herbivorous insects. The combined impact of drought stress and insect herbivory on plant metabolism has rarely been studied, particularly in woody plants. In this study, we investigated the influence of varying degrees of drought, both alone and in combination with herbivory by the leaf beetle Chrysomela tremulae, on the morphological and chemical characteristics of black poplar (Populus nigra) trees using a full factorial experimental design. We quantified morphological traits, volatile organic compound (VOC) emissions, phytohormone and amino acid concentrations, and phenolic profiles. Drought conditions increased the concentrations of salicylic acid (SA) and abscisic acid (ABA), while feeding induced ABA and SA. Amino acid profiles shifted significantly under drought conditions, particularly in beetle-infested plants. In contrast, salicinoids, which are the most important phenolic defense compounds in poplars, remained relatively stable. We also observed significant compound-specific effects on both constitutive and herbivore-induced VOC emissions. Our results demonstrate that drought and insect herbivory exert a joint influence on the chemical responses of P. nigra across multiple metabolic pathways. These findings highlight how the interaction between abiotic and biotic stresses can influence the defense chemistry of trees, which will consequently affect ecological interactions in forest ecosystems in the face of climate change.

Maple syrup production is strongly influenced by spring weather conditions, particularly the frequency and intensity of freeze-thaw cycles. However, marked individual differences in sap and sugar yields persist among trees growing under similar stand conditions, indicating additional tree-level sources of variation. This study aimed to explain inter-individual variability in maple yields in commercial high-vacuum syrup production using structural, morphological, and growth characteristics. We used terrestrial light detection and ranging to derive variables describing crown, stem, and whole-tree size, biomass, and structure in 38 mature sugar maples. We related these variables to individual yields, including sap volume, sugar content, and syrup production. Our models explained substantial inter-individual variability: 52% in sap sugar content, 44% in sap volume, and 47% in syrup production. Laterally expanded crowns were associated with higher sap sugar content, as were lower growth rates in the first 25 mm of wood. Sap volume was highest in large, heavily branched trees with crowns extending vertically along the stem. Projected crown surface area was the strongest predictor of syrup yield, with an estimated increase of 250 mL per 5.6 m2. These findings highlight the importance of multidimensional crown development in maximizing individual yields in maple syrup production.

Heat tolerance determines the vitality of tree species under climate change independently of drought tolerance, but has been much less studied than tree water relations. We studied species-specific differences and the capacity for seasonal heat acclimation in Central Europes naturally most important tree species, Fagus sylvatica, in comparison with two exotic tree species (Fagus orientalis, Pseudotsuga menziesii) that are considered for silvicultural climate change adaptation in managed forests. Foliage of mature trees was incubated at temperatures from 35-50 {degrees}C for up to 4 h to simulate daily heat maxima during heat waves. The maximum quantum yield (Fv/Fm) of photosystem II (PS II) of dark-adapted leaves was measured, because the PS II is particularly sensitive to heat and its functionality can decide on plant survival under heat. Fagus sylvatica was much more tolerant to heat than Pseudotsuga menziesii, but weakly (albeit significantly) less tolerant than Fagus orientalis. Within its limits, Pseudotsuga menziesii showed high seasonal heat acclimation with constantly increasing tolerance during the growing season. Fagus orientalis, but practically not Fagus sylvatica, also acclimated to heat. This makes Fagus orientalis slightly superior over Fagus sylvatica in terms of heat tolerance, whereas the suitability of Pseudotsuga menziesii for silvicultural climate change adaptation is questionable. Strong heat acclimation, but also overall low heat tolerance, in Pseudotsuga menziesii might be the result of evergreenness, which requires the generation of both cold and heat tolerance during the year.

O_LIEuropean beech (Fagus sylvatica L.) is a widely distributed, ecologically and economically important deciduous tree species in European forests, but is increasingly threatened by drought stress. Volatile organic compounds (VOCs) are ubiquitous plant metabolites that may serve as non-invasive biomarkers of drought stress, yet they have rarely been studied in European beech. C_LIO_LIIn this study, we examined VOC responses of European beech to experimental drought across diverse genetic backgrounds in a common garden. The 72 four-year-old beech saplings represented three genetic clusters, seven provenances (geographic seed sources), and 12 maternal seed families. Half of the saplings were assigned to the drought treatment and received no water for 14 days, while the remaining saplings served as controls and were watered as required. VOC profiles, quantified as peak heights of mass spectral features, were measured for all individuals during pre-drought, drought, and rewatering periods. C_LIO_LIWe found that pre-drought VOC profiles, in particular monoterpenes, varied significantly among genetic backgrounds. Experimental drought significantly altered VOC profiles, characterized by increased green leaf volatiles and decreased monoterpenes, oxidized terpenoid derivatives, and other fatty acid derivatives. Reductions in monoterpenes persisted after rewatering, indicating a drought legacy effect. Drought responses were largely conserved across genetic backgrounds, with significant seed family-specific responses detected for only three VOC features. C_LIO_LIOur findings suggest that VOC profiles are genetically structured yet highly plastic under drought and highlight their potential as non-invasive biomarkers for monitoring drought stress in European beech under climate change. C_LI

Subalpine forests in the Alps are fragile ecosystems that play a crucial role in regional water resources and the local climate. These ecosystems are ecologically significant due to their unique biodiversity and vulnerability to climate change. While several components of the hydrological balance have been studied, the interplay between catchment-scale processes and plot-scale drivers such as fog presence and forest age remains insufficiently understood. To address this, we investigated the hydrological balance of a subalpine coniferous forest catchment at the Renon site in the Italian Alps, integrating observations across spatial scales. The study area includes a mosaic of mature and younger regrowth forest, where both interannual and seasonal variability in precipitation and fog presence are pronounced. At the catchment scale, we quantified above-canopy precipitation, evapotranspiration (ET, measured via eddy covariance at the ICOS tower), stream discharge, and soil moisture dynamics. Within the catchment, we characterised water partitioning using sap flow sensors for tree transpiration, throughfall and stemflow collectors with rain gauges above and below the canopy and epiphyte sampling. Mixed fog-rain events frequently coincided with higher throughfall. However, these changes had a minor effect on soil water storage and catchment discharge in the annual water balance, which was nearly closed. At the plot scale, our results show that tree transpiration was higher in the younger forest structure, while canopy interception is a dominant process in water partitioning in the older forest structure, where lichen abundance likely enhances interception. This study highlights the importance of multi-scale monitoring in temperate mountain forests, where forest age influences water partitioning, and fog presence, though not directly quantified, can still contribute to reducing evaporative processes. Such contributions may gain importance under changing climate conditions, albeit less prominently than in tropical or subtropical cloud forests.

O_LIIntraspecific variability in hydraulic and morphological traits may alter projections of forest vulnerability to climate change. Recognizing multiscale nature is essential, as trait relationships within populations may not reflect those across the species range. C_LIO_LIWe assessed variability in 11 traits in 10 natural populations of Abies alba across environmental and phylogeographic gradients to evaluate potential impacts on drought vulnerability. We quantified trait variation within and among populations, tested effects of climate, phylogeny and local factors, and evaluated trait coordination across scales. C_LIO_LIHydraulic safety traits and wood density showed low variability, indicating strong constraints. In contrast, water-use and efficiency traits were highly variable. Aridity influenced several traits, but reduced variance was detected only for leaf residual conductance and succulence consisted with stabilising selection. Trait coordination was weak within populations than among populations. C_LIO_LIOverall, A. alba combines constrained hydraulic safety traits with variable water use traits. This may buffer drought impacts but limits shifts in hydraulic safety margins, potentially increasing risks of hydraulic failure, especially in humid environments where populations lack the capacity to tolerate prolonged dry periods. Thus, our findings highlight the need to account for scale-dependent trait variability, coordination, and their underlying drivers when predicting species adaptive capacity to drought. C_LI

SummaryO_LIIn woody perennials, reproductive allocation considered as a fraction of NPP (RA) has been rarely quantified, especially tracked across plant lifetimes, measured in biologically meaningful currencies, and separated from standing biomass. C_LIO_LIWe addressed this gap by measuring dry mass, nitrogen, and phosphorus for every aboveground tissue type for 14 iteroparous perennial shrubs tracked across their full lifetimes, calculating RA using four accounting schemes and three currencies. C_LIO_LIRA was substantially higher by mid-life than typical estimates from ecosystem-scale studies. Distinguishing standing biomass from yearly production further revised RA upwards. Using a nutrient currency (especially P) increased the perceived costs of reproduction. Propagules were highly nutrient-enriched and reproductive accessory costs consumed more nutrients than did the propagules themselves. Considering N and P resorption from senescing leaves and wood shrank the effective vegetative nutrient budget, further concentrating net annual nutrient demand in reproductive tissues. C_LIO_LIOur results highlight high investment in RA for woody perennials, especially using nutrient currencies. Broadly similar allocation patterns were observed across species with different functional traits and lifespans, suggesting generality that may apply across biomes. Widespread underestimation of RA in forest growth models likely overestimates the proportion of NPP available for vegetative growth, leading to substantial errors in predictions. C_LI

Winter warming is altering plant exposure to cold events, yet its effects on seasonal cold hardiness dynamics remain poorly understood. Here we quantified bud cold hardiness across four dormant seasons in a boreal peatland forest whole ecosystem warming experiment. Across a +0.00 to +9.00{degrees}C warming gradient, we semi-regularly measured cold hardiness in two overstory (Larix laricina and Picea mariana) and two understory species (Chamaedaphne calyculata and Rhododendron groenlandicum). Warming reduced cold hardiness in fall and spring by delaying acclimation and advancing deacclimation. However, risk was only increased in late winter and spring for three species. Warming reduced snow cover, increasing temperature variability and cold damage to understory shrubs. Together, our results show that cold damage risk depends on species traits, microclimate, and seasonal timing.

Internal decay in living trees is an important component of carbon and nutrient cycling as well as species and structural diversity maintenance in forest ecosystems. We used sonic and electrical resistance tomography to evaluate and compare the prevalence and severity of stem decay in 57 living trees among four common species (Acer rubrum L., Nyssa sylvatica Marsh., Quercus rubra L., and Tsuga canadensis (L.) Carriere)) with overlapping and non-overlapping distributions across wetland and upland habitat types at the Harvard Forest in Petersham, MA, USA. Independent of tree size, site identity best explained variation in the prevalence of decay across trees sampled, whereas species identity best explained the severity of decay. We categorized trees as having no decay, incipient decay, active decay, or cavities based on combined sonic and electrical resistance metrics, the latter generated by a custom image analysis application. About 31% of wetland trees exhibited incipient decay (compared to 11% in the upland), whereas about 32% of upland trees exhibited active decay (compared to 10% in the wetland). Our study highlights a new quantitative framework for decay categorization through normalized principal component analysis (PCA) and decay analysis software that complements dual tomographic methodology for future investigations of ecological drivers of decay presence and susceptibility.

Forest regeneration depends not only on how many seeds trees produce, but on the physiological quality of those seeds. Yet while climate-driven shifts in seed quantity and masting have received sustained attention, the parallel question of whether climate change degrades seed quality remains poorly resolved. Using a nationwide dataset of seed mass and viability in European beech (Fagus sylvatica L.) collected between 1996 and 2024 (13,349 seed lots from 381 forest districts across Poland), with climate-quality analyses focused on 5,374 freshly harvested seed lots from 353 districts (2004-2023), we asked whether the two components of seed quality respond to different seasonal climatic windows, and whether harvest-year climate also shapes seed performance during long-term cold storage. Seed mass and seed viability were only weakly correlated (Spearmans {rho} = 0.15), acting as two independent dimensions of seed quality. Both revealed substantial temporal variation over the study period, but along distinct trajectories. Seed mass declined markedly between segmented-regression breakpoints in 2009 and 2019, more steeply at higher latitudes, coinciding spatially and temporally with the masting breakdown reported at the species northeastern range margin. Climatic associations were correspondingly divergent. Viability was positively associated with previous summer temperature, consistent with temperature-cued flower initiation, and negatively with spring temperature in the harvest year, plausibly reflecting thermal disruption of early embryogenesis. Seed mass showed no significant association with any seasonal climatic predictor, indicating control by slower or unmeasured processes. Storage duration progressively reduced viability, and this decline was further modulated by climate during seed development, with seeds developing under climatically suboptimal conditions losing viability faster. These results expose a hidden decoupling between seed quantity and seed quality under contemporary climate change, with direct consequences for forest regeneration and for ex situ conservation strategies that assume mast-year seeds will remain viable for decades.

Norway spruce (Picea abies) responds to attacks by the spruce bark beetle (Ips typographus) through the rapid activation of local defense mechanisms, but field studies can be difficult to standardize due to variable attack pressure and environmental heterogeneity. Here, we developed a phytotron-based assay that mimics early beetle-associated stress using insect-derived protein extracts, enabling reproducible molecular analyses under controlled conditions. Ten-week-old spruce seedlings were stem-treated with mock buffer or beetle protein extracts, followed by transcriptomic analyses of stem tissues and targeted metabolomic profiling of needles at 2 and 48 h post-inoculation. RT-qPCR analysis revealed rapid transcriptional activation of signaling and defense genes in Norway spruce, with NP-40-based protein extracts producing the most consistent early response. RNA-seq analysis revealed transcriptional dynamics, with 488 differentially expressed genes detected at 2 h and 84 at 48 h post-inoculation relative to mock-treated controls. Early responses at 2 h were characterized by activation of genes associated with immune perception and signal transduction. By 48 h, the response shifted toward accumulation of transcripts encoding defense proteins such as chitinases, defensins, proteinase inhibitors, and pathogenesis-related (PR) proteins. Importantly, a substantial proportion of differentially expressed genes overlapped with those previously identified in mature Norway spruce trees during pioneer bark beetle attack under field conditions, supporting the biological relevance of the assay. In contrast, targeted analyses of secondary metabolites performed in needle tissue revealed limited systemic changes across time points, suggesting that early induced defenses may remain largely localized to the stem. Together, these results demonstrate that beetle-derived proteins trigger a rapid and temporally structured defense response in Norway spruce seedlings and establish a reproducible elicitor-based platform for dissecting conifer immune responses and screening spruce genotypes for bark beetle resistance. HighlightBark beetle protein elicitors trigger temporally structured immune responses in Norway spruce seedlings that overlap with responses observed in mature trees, with rapid immune signaling at 2 h followed by defense protein accumulation at 48 h.

The composition and quantity of root exudates are strongly influenced by physiological and environmental conditions, reflecting dynamic changes in plant metabolism. Although many studies report that root exudate metabolite profiles vary with plant phenology, few have disentangled the effects of phenological stage from those of nutrient availability. We collected root exudates from white lupin (Lupinus albus) grown in a fine phosphorus (P) gradient (5, 10, 20, 30 and 50 {micro}M P) in hydroponics at three developmental stages, performed untargeted metabolomics using GC-MS, and measured 10 above and belowground traits. Our results show that plant phenological stage exerts a stronger influence on exudate metabolomic profiles than variation in P supply. During leaf development, exudates were dominated by metabolites associated with carbon metabolism, whereas flowering was characterized by compounds related to secondary metabolism and cell wall turnover. Phosphorus influenced exudate profiles only at the flowering stage, with distinct profiles observed at 5 and 10 {micro}M P compared with 20-50 {micro}M P. These findings provide new insights into the temporal regulation of root exudation and demonstrate that plant developmental stage is a primary determinant of metabolic responses to phosphorus availability and, potentially, rhizosphere functioning under nutrient limitation. HighlightRoot exudate quantity and composition in white lupin is shaped more by plant phenological stage than phosphorus supply, with phosphorus effects emerging only during flowering.

Large old trees are of eminent importance for organic carbon storage in forest ecosystems and thus play a role in mitigating climate change. Such trees also have an increased risk of internal stem decay and tree cavity formation, which promotes biodiversity, but complicates the prediction of their biomass and carbon stocks, which is usually done from stem diameter and tree height data applying allometric biomass functions. Since the extent of internal stem decay is known to vary widely between different forest ecosystems and data from moist temperate forests exhibited low significance of internal stem decay, we studied dry, frequently fire-exposed Pinus ponderosa forests in central Oregon to capture the other climatic extreme of temperate forests. We hypothesized high significance of internal stem decay for stand aboveground tree biomass, as we assumed widespread stem injury from fire. In addition, we tested the hypothesis that far more than the largest 1% of trees are necessary for 50% stand biomass, as this hypothesis is found in the literature, but has been challenged in other studies. We found low biomass loss due to internal stem decay by only ca. 1% suggesting that also for fire-prone temperate forests of western North America, biomass estimates based on allometric regression are reliable. The 1% largest trees-50% stand aboveground biomass hypothesis has to be rejection for our forests as long as only trees of a size are included that noteworthily contribute to stand biomass. This metrics strongly depends on regeneration density, which is not relevant for stand biomass.