Tree Physiology

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.

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.

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.

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

Grasslands are vulnerable to increasing drought with global warming, but process-based models lack the mechanistic knowledge required to predict the magnitude of drought impacts. While a plant hydraulics framework has been successful in advancing process understanding of drought responses in trees, and how drought responses vary across rainfall gradients, similar approaches have rarely been applied to grasses. Here, we quantified the progression of key drought response processes in sixteen dominant perennial grasses (seven C3 and nine C4) with differing climatic origins across eastern Australia. We found that stomatal closure, hydraulic impairment and leaf browning occurred concurrently, in contrast to the progressive sequence typically observed in trees. We also found that drought response traits were not correlated with species climate of origin. The early impairment of leaf hydraulic conductance and leaf browning along with the lack of correlation with climate of origin suggest that grasses may employ fundamentally different strategies to adapt to low water availability than trees. These results highlight the need for grass-specific parameterization of drought responses in process-based models.

Transgenerational plasticity can shape plant phenotype and influence plant response to environmental changes in interaction with the current conditions. While how past stress interact with either current optimal or stress conditions is increasingly documented within a single plant, transgenerational plasticity remains particularly poorly understood especially at the metabolome level. In our study, we investigated whether heat stress induces transgenerational metabolic and phenotypic modifications along two successive clonal ramet generations of the sub-Antarctic aquatic plant Limosella australis. We performed untargeted metabolomic approaches and measured morphologic and performance traits, to assess both transgenerational plasticity of the metabolome and the phenotype. We found that heat stress remodelled the metabolic profile and influenced the foraging strategy of our clonal plant, and that some of these metabolic changes persisted into the first clonal generation. This one therefore adopted an intermediate growth strategy, even though culture conditions were optimal. By comparing differentially accumulated features between daughter ramets from heat stressed mother ramets and from unstressed mother ramets, we identified common and specific metabolites accumulation to heat stress response, belonging to diverse compound families. However, we did not observe any adaptative advantage and any metabolic imprint during another heat stress applied on the second clonal generation. This work provides especially new clues into how plant metabolome integrates and transfers previous stressed clonal generation's information.

As extreme heat events increase in frequency and intensity worldwide, understanding how woody perennial crops respond to higher maximum temperatures is critical. Perennials face distinct challenges, persisting across many seasons under increasingly variable and extreme conditions, and heat waves threaten the viability of wine grape cultivars through impacts on yield, wine quality, and long-term vine health. To test whether irrigation practices before and during heat waves affect grapevine carbon (C) storage and health, we experimentally manipulated irrigation regimes surrounding heat waves from 2019-2021 in a commercial Cabernet Sauvignon vineyard in the Lodi AVA of Californias Central Valley. Vine physiological traits and yield were measured throughout, and whole-vine nonstructural carbohydrate (NSC) concentrations were quantified after three growing seasons. Although lower supplemental irrigation reduced photosynthesis, stomatal conductance, and fruit yield, whole-vine NSCs did not differ significantly in any perennial organ by the experiments end, indicating that reproductive output and final NSC status responded to irrigation on different timescales. These results suggest that moderate supplemental irrigation during heat events is sufficient to mitigate negative impacts on yield and quality while supporting recovery of NSC reserves, though longer-term monitoring is needed to confirm that this short-term resilience persists.

The xerophyte argan (Sideroxylon spinosum) has great ecological and socioeconomic importance for Morocco. However, it is endangered due to climate change and human overexploitation. We assess drought resistance traits of argan and test the potential of arbuscular mycorrhizal (AM) symbiosis to promote its growth and mitigate the consequences of drought. We compare ten endemic Moroccan mycorrhizal inocula with the model AM fungus Rhizophagus irregularis and with the drought-adapted isolate Diversispora omaniana. We integrated physiological phenotyping and RNA sequencing to investigate the stress resistance mechanisms of argan against drought. We show that AM symbiosis significantly mitigates drought effects on plant growth, mainly by improving water relations and photosynthetic efficiency, resulting in increased growth rates. Taken together, physiological and transcriptomic analyses show that stress markers were moderatly induced during severe drought stress irrespective of mycorrhizal status, suggesting that argan adopts a drought-coping strategy that involves both, stress avoidance and stress tolerance. Argan is highly AM-responsive, both at the phenotypic and transcriptomic level, suggesting that AM has great potential to promote argan growth under drought stress.

Background and AimsThe development of unisexual flowers relies on the tight coordination of flower organ identity and sex determination. The genus Quercus is typically considered strictly monoecious, bearing fully segregated male and female flowers within the same individual tree. However, several reports of atypical flowering across the genus challenge this canonical view, suggesting that flowering in oaks may be more flexible than traditionally assumed. In this work, the dynamics of flower development in Quercus orocantabrica were examined to correlate contrasting floral morphologies with divergent molecular profiles. MethodsThe flowering phenology of Q. orocantabrica trees was closely monitored over several individuals and years, together with a detailed floral morphological analysis of male, female and atypical flowers. Key floral homeotic gene homologues were identified, and their expression assayed in the development of different flowers. Key ResultsRecurrent and widespread hermaphroditic flowering was detected in several Q. orocantabrica trees, frequently associated with unseasonal flowering events. Gene expression analysis of male, female and hermaphroditic flowers revealed a sex-biased expression of Q. orocantabrica B- and C-class genes, with the B-class gene QoPI in particular being tightly associated with the presence of fully-developed stamens. In addition, the expression of the C-class gene QoSHP contrasted with reports in other Fagaceae, highlighting a potential functional divergence of the C/D-class lineage within the family. ConclusionsThe results here depicted indicate that the dynamics of floral sex identity in oaks are more plastic than traditionally assumed, supporting a reinterpretation of oak reproductive biology based on a versatile and resilient framework responsive to different developmental contexts.

Drought is a major abiotic stress threatening plant productivity and agricultural sustainability, yet the molecular mechanisms underlying adaptive root responses to water deficit in the water use strategies continuum remain insufficiently understood, particularly in perennial crops. In this study, we explored drought responses in nine accessions belonging to three wild Vitis species (V. acerifolia, V. candicans, and V. doaniana) displaying varying drought-response strategies. Plants were subjected to moderate drought stress (40% soil water content) for three weeks under greenhouse conditions. By integrating physiological, metabolic, and transcriptomic analyses, we aimed to identify both conserved and species-specific mechanisms associated with drought adaptation. Differential expression analyses revealed a conserved core set of drought-responsive genes shared among species, including genes involved in abscisic acid signaling, reactive oxygen species detoxification, solute transport, and plant defense. In parallel, each species exhibited distinct transcriptional and metabolic signatures reflecting alternative adaptive strategies related to osmoregulation, and oxidative stress mitigation. Weighted gene co-expression network analysis (WGCNA) further revealed significant associations between constitutive, non-plastic gene expression and root phenotypic traits. Overall, our findings demonstrate that wild Vitis species rely on both conserved stress-responsive pathways and species-specific constitutive regulation to cope with drought stress. These results highlight the importance of root-associated traits and intrinsic regulatory networks in shaping drought adaptation and provide new targets for the development of drought-resilient grapevine rootstocks.

Tree early life stages are particularly sensitive to warming, yet their responses remain poorly understood despite their importance for forest regeneration. Here, we investigated how warming affects early-life traits in two widespread European white oaks: Quercus pubescens and Q. petraea. We conducted a common garden experiment using 17 populations exposed to three temperature regimes. We measured 19 traits encompassing germination, phenology, and functional and fitness-related traits and performed individual trait mixed-effects models based on temperature transfer distance and the climate of the population. We found that population climate was the primary driver of early stages traits responses to warming, with climatic drivers varying strongly among traits and species. Particularly in Q. pubescens, warmer and drier populations showed lower fitness (germination and survival percentages, total biomass) that declined further under warming, consistent with a cost of drought avoidance strategies under continuously wet conditions; in Q. petraea, continental populations outperformed others at low temperature transfer distance but suffered the steepest fitness declines under further warming, suggesting a narrow thermal optimum shaped by cold adaptation. Warming generally advanced germination and leaf emergence, increased leaf pigment concentrations and fine-root allocation, reduced specific leaf area. Extreme warming reduced survival, growth and germination. Nevertheless, moderate warming (+0 to +5{degrees}C) was rarely detrimental and sometimes beneficial. Our results demonstrate that population climatic origin is a key determinant of regeneration responses to warming, highlighting the need to consider within-species adaptive variation to understand forest regeneration potential under climate change.

Rising winter temperatures threaten deciduous fruit tree productivity by depleting carbohydrate reserves during dormancy. This study investigated Stem Photosynthetic Capacity (SPC), a rare adaptive trait from wild Prunus arabica, as a mechanism to enhance almond carbon economy. Using extreme segregating groups from the F1 population (P. dulcis X P. arabica), we evaluated physiological performance through high-resolution lysimetric and multi-year orchard monitoring. High-SPC [SPC(+)] genotypes maintained significantly greater stem CO2 assimilation and transpiration during leafless periods compared to low-SPC [SPC(-)] progenies. Over five successive seasons, SPC(+) trees exhibited a 33.3% increase in trunk secondary growth and reached 10% bloom approximately 8 days earlier. Most importantly, the SPC(+) group achieved a 4.6-fold increase in mean kernel yield when compared to SPC(-) group. These findings demonstrate that SPC provides a flexible, supplementary winter carbon source that directly supports both vegetative and reproductive development. Integrating SPC into commercial almond breeding programs may offer a valuable strategy to improve climate resilience and help sustain yields under warming conditions. HighlightIntegrating stem photosynthesis into commercial almond hybrids provides a winter carbon source that advances blooming, expands trunk growth by [~]33%, and increases kernel yields by more than 4.5-fold.

Global climate change has increased the frequency and intensity of drought events, making it urgent to understand how native species respond to water deficit (WD). In biodiverse environments such as tropical forests, simple methods are needed to study multiple species simultaneously. This can help predict how natural environments will respond to climate change and guide the strategic selection of drought-resistant species for reforestation. This study aimed to: (1) adapt an existing simple and inexpensive method to apply a controlled WD on tree seedlings from tropical species commonly produced in nurseries for restoration projects, suitable for greenhouse experiments; and (2) evaluate the effectiveness of this method in generating ecophysiological responses to WD that allow the estimation of species' drought resistance. Ten native tree species from the Semideciduous Seasonal Forest (SSF), a phytophysiognomy of the Atlantic Forest, were selected. An existing method was adapted to implement capillary irrigation, in which the bases of the seedling tubes were placed in floral foam blocks positioned inside 15 L plastic containers filled with water. A gradual and severe WD was applied to five seedlings of each species by removing all water from the containers, leaving only the water retained in the saturated floral foam available for plant uptake. The remaining seedlings were maintained well-watered (containers full and foam saturated) as the control group. Stomatal conductance (gs) was measured daily for all seedlings until they reached 50% or less of their initial gs (igs); at this point, stem water potential ({Psi}w) was measured. Both gs and {Psi}w differed significantly among treatments and species (p < 0.01). Ficus guaranitica and Heliocarpus popayanensis were the only species that did not show significant {Psi}w differences between treatments, indicating higher drought resistance. In contrast, Campomanesia xanthocarpa and Eugenia uniflora had the lowest {Psi}w values under WD, suggesting lower drought resistance. The remaining species were distributed along a gradient of responses to WD. Additionally, no correlation was found between {Psi}w and gs at 50% igs in the WD group (rho = 0.16, p = 0.26). The method proved effective in inducing controlled WD and generating measurable ecophysiological responses, offering a useful tool for screening native species for drought resistance.

Direct heat damage has been considered secondary as a cause of climate change-induced tree mortality and productivity declines in forests compared with climate change effects on tree water relations. However, evidence from temperate and tropical forests is accumulating that direct heat damage in the photosystem II (PS II) that is independent of water relations is also a realistic scenario under climate change. We analyzed PS II heat tolerance in Larix sibirica, which represents a dominant boreal tree species in Siberia and northern Central Asia in cold environments with subzero or near-zero mean annual temperatures, but nevertheless warm summers. Thermal imaging was applied to relate heat thresholds found in the laboratory to canopy temperatures in forests on north-facing mountain slopes, which are the main habitat of L. sibirica. L. sibirica showed slight decreases of the maximum quantum yield of PS II (Fv/Fm) at 35{degrees}C and 40{degrees}C after up to 4 h, but strong reductions at [&ge;]45{degrees}C and minor increases in Fv/Fm in late summer, which could be interpretation as heat acclimation. Canopy temperatures in the study year did not reach the thresholds for serious PS II heat damage. However, L. sibirica was more strongly sensitive to heat than temperate conifers. This first combined study of heat tolerance and canopy temperatures from boreal forests points to the possibility of low heat tolerance of boreal tree species, but such conclusion would require the study of more tree species.

O_LITerrestrial biosphere models commonly use empirical scaling factors to represent soil moisture constraints on carbon and water fluxes, but these lack mechanistic grounding and produce inconsistent estimates of soil moisture limitations on primary production and transpiration across models. C_LIO_LIHere we extended the least-cost hypothesis for optimal stomatal conductance to account for soil moisture limitations by allowing soil water availability to modulate the carbon cost of water transport, drawing on the observed temperature dependence of stem respiration, and derived a simple empirical approximation to the theory using global {delta}13C and eddy covariance data. C_LIO_LIThe empirical analysis shows moderated thermal acclimation of stem respiration and a weak increase in water transport costs with aridity, supporting the interpretation that the decline in light-use efficiency (LUE) under arid conditions is primarily attributable to non-stomatal limitations. C_LIO_LIValidation against an independent global dataset of sapflow-derived canopy conductance and transpiration shows that the revised scheme significantly improves the predictive power of the least-cost hypothesis, offering a more mechanistically coherent alternative to existing soil moisture parameterisations. C_LI

Biuret, a nitrogen-rich by-product of urea and a common contaminant of urea-based fertilisers, has long been considered a passive phytotoxin, affecting plant performances. Yet its effects on root development and the existence of endogenous mechanisms of perception or tolerance remain largely uncharacterised. Here we combined physiological, developmental, genetic and transcriptomic approaches to investigate the response of Arabidopsis thaliana to biuret. Biuret inhibited primary root growth in a dose-dependent manner by reducing meristematic cell division rather than cell elongation, and concomitantly impaired shoot growth by limiting leaf expansion. This root inhibition was reversible upon biuret removal and was accompanied by increased auxin-responsive (DR5) and decreased cytokinin-responsive (TCS) outputs at the root apex, consistent with a regulated remodelling of meristem activity rather than purely cumulative damage. A forward genetic screen identified the biuret-resistant mutant bir29, which sustained root and inflorescence development under inhibitory concentrations. Using {superscript 1}N-labelled biuret, we showed that resistance occurred without any change in biuret influx or accumulation, uncoupling sensitivity from exposure. Whole-genome transcriptomics revealed that bir29 fails to execute the wild-type response, neither repressing the cell-cycle machinery nor deploying the stress-associated programme induced by biuret. Genetic characterisation linked resistance to multiple genomic loci required for full resistance. Together, the results indicate that biuret triggers an active, reversible and genetically tractable developmental response, suggesting that this xenobiotic compound is integrated into endogenous signalling networks. Significance StatementBiuret, a poorly metabolised contaminant of urea fertilisers, is generally regarded as a passive phytotoxin, yet we show that it inhibits Arabidopsis root growth through a reversible and genetically tractable developmental response, accompanied by reorganised auxin and cytokinin signalling, rather than through cumulative chemical injury. The isolation of the resistant mutant bir29 suggests that plants integrate this xenobiotic molecule into endogenous signalling networks, reframing biuret as an informative probe of root developmental regulation.

Background and AimsPaspalum notatum reproduces through either sexuality or apomixis, two pathways that may coexist within the same individual and are regulated by interconnected molecular networks responsive to environmental cues. Here, we characterized the transcript structure and expression of BUD SITE SELECTION PROTEIN 13 (BUD13), a component of the RES spliceosomal complex previously reported as differentially expressed in florets of sexual and apomictic plants, as a first step toward testing its involvement in the molecular regulation of the apomixis-sexuality switch. MethodsPreviously generated floral and leaf transcriptomes from sexual and apomictic Paspalum notatum plants, including Oxford Nanopore long-read data, were mined to characterize BUD13 transcript structure and expression. Phylogenetic analyses and in silico mapping were conducted to infer evolutionary relationships and determine the origin of the transcripts. Differential expression was validated by RT-qPCR, while in situ hybridization was used to reveal cell-specific ovule expression patterns. Key resultsBUD13 is expressed in Paspalum notatum florets as a truncated isoform (SHORT) encoding a small protein lacking part of the herpes simplex virus regulatory protein (ICP4) domain. Two SHORT transcripts, SHORT1 and SHORT2, with different 5' untranslated region (UTR) regions, were identified in flowers. SHORT1 was consistently upregulated in apomictic ovules from premeiosis to anthesis. Both transcripts originated from a single genomic locus located in the subtelomeric region of the short arm of chromosome 6. SHORT isoforms with variable structures were detected in other monocots. In situ hybridization showed that, whereas BUD13 was expressed throughout sexual ovules, expression was absent from the female germline of apomictic ovules. A consistent expression was observed in somatic proembryos of aposporous embryo sacs. ConclusionsOur findings reveal structural, spatial and temporal divergence in BUD13 expression between sexual and apomictic reproductive programs, providing new insights into the molecular regulation of asexual seed formation.

Benzoxazinoids are a paradigmatic class of indole-derived specialized metabolites released into the soil through root exudates and originally studied for their allelopathic and toxic effects on neighboring plants, herbivores, and microorganisms. They are now recognized as regulators of diverse plant-organism interactions, with beneficial effects such as in microbiome-mediated resistance to pathogens in plant successions. However, the mechanisms by which benzoxazinoid-containing root exudates contribute to pathogen control beyond microbiome structuring remain unclear. Here, using an agriculturally relevant rice-maize co-culture system, we show that benzoxazinoids naturally exuded by maize roots are taken up by rice roots and are associated with reduced rice blast disease in leaves. This protection occurs without detectable benzoxazinoids accumulation, constitutive immune activation, or growth penalty in rice leaves. Instead, maize-derived benzoxazinoids uptake in rice roots is associated with chromatin hyperacetylation at, and increased expression of key phenylpropanoid biosynthetic genes, and broad metabolome reconfiguration. These effects extend systemically to leaves, where rice establishes a defense-related chemical state distinct from systemic acquired resistance as observed in benzoxazinoid-dependent, microbiome-mediated plant-soil feedbacks. Our findings support a model in which specialized metabolites released through root exudation by one crop species can be acquired by a neighboring species and trigger chromatin-associated metabolic reprogramming linked to systemic chemical defenses. This work provides a molecular framework connecting plant-plant interaction, root exudates, chromatin regulation, systemic chemical defense, and disease suceptibility, opening new perspectives for exploiting natural plant-plant chemical interactions in sustainable and resilient agroecosystems. Significance statementHormesis, whereby low or moderate exposure to otherwise harmful compounds stimulates adaptive beneficial responses, has rarely been used as a conceptual framework for understanding interactions between organisms. Here, we show that naturally exuded benzoxazinoids, specialized metabolites historically viewed mainly as broadly toxic coumpounds in exogenous treatments, can trigger beneficial systemic responses in a neighboring crop species. In rice, root acquisition of maize-derived benzoxazinoids is associated with histone hyperacetylation, defense-related metabolome reprogramming, and reduced susceptibility to rice blast disease in leaves without obvious growth penalty. These findings reveal a hormesis-like process in plant-plant interactions, in which interspecific chemical exposure modulates receiver physiology rather than simply causing toxicity. More broadly, this work suggests that beneficial responses to natural-dose chemical cocktails may be an underexplored driver of adaptive responses in complex biological systems, with implications for agroecology, ecotoxicology, and exposome research.

The Eurasian spruce bark beetle Ips typographus, a major forest pest on Norway spruce (Picea abies), forms intimate associations with several types of microbial symbionts. While previous research has focused primarily on filamentous fungi, yeasts have remained largely unexplored. Here, we show that yeasts associated with I. typographus may contribute to host tree colonization by providing defensive benefits. Dominant yeasts (Yamadazyma, Kuraishia, Nakazawaea, and Wickerhamomyces), which are phylogenetically related to other insect-associated Saccharomycotina, significantly attract adult beetles. Moreover, several yeasts inhibit the growth of the pathogenic fungus Trichoderma harzianum in vitro, and beetle eggs benefit from the presence of Kuraishia capsulata by reduced fungal infection under semi-natural conditions. Strikingly, these effects are mediated by the yeasts' transformation of the tree's defensive stilbene glycosides into antimicrobial aglycones and phenolic acids that accumulate in beetle galleries. These findings reveal a previously unrecognized role of symbiotic yeasts in converting spruce defensive stilbene glycosides into antimicrobial aglycones and oxidative cleavage products that accumulate in beetle galleries, and might contribute to the survival of their bark beetle host.