Ancient trees are essential elements for high-mountain forest conservation: Linking the longevity of trees to their ecological function.

Mature forests and their extremely old trees are rare and threatened ancient vestiges in remote European high-mountain regions. Here, we analyze the role that extremely long-living trees have in mature forests biodiversity in relation to their singular traits underlying longevity. Tree size and age determine relative growth rates, bud abortion, and the water status of long-living trees. The oldest trees suffer indefectible age-related constraints but possess singular evolutionary traits defined by fitness adaptation, modular autonomy, and a resilient metabolism that allow them to have irreplaceable roles in the ecosystem as biodiversity anchors of vulnerable lichen species like Letharia vulpina. We suggest that the role of ancient trees as unique biodiversity reservoirs is linked to their singular physiological traits associated with longevity. The set of evolutionarily plastic tools that can only be provided by centuries or millennia of longevity helps the oldest trees of mature forests drive singular ecological relationships that are irreplaceable and necessary for ecosystem dynamics.

Abscisic acid triggers vitamin E accumulation by transient transcript activation of VTE5 and VTE6 in sweet cherry fruits.

Tocopherols are lipophilic antioxidants known as vitamin E and synthesized from the condensation of two metabolic pathways leading to the formation of homogentisate and phytyl diphosphate. While homogentisate is derived from tyrosine metabolism, phytyl diphosphate may be formed from geranylgeranyl diphosphate or phytol recycling from chlorophyll degradation. Here, we hypothesized that abscisic acid (ABA) could induce tocopherol biosynthesis in sweet cherries by modifying the expression of genes involved in vitamin E biosynthesis, including those from the phytol recycling pathway. Hence, the expression of key tocopherol biosynthesis genes was determined together with vitamin E and chlorophyll contents during the natural development of sweet cherries on the tree. Moreover, the effects of exogenously applied ABA on the expression of key tocopherol biosynthesis genes were also investigated during on-tree fruit development, and tocopherols and chlorophylls contents were analyzed. Results showed that the expression of tocopherol biosynthesis genes, including VTE5, VTE6, HPPD and HPT showed contrasting patterns of variation, but in all cases, increased by 2- and 3-fold over time during fruit de-greening. This was not the case for GGDR and VTE4, the first showing constitutive expression during fruit development and the second with marked down-regulation at ripening onset. Furthermore, exogenous ABA stimulated the production of both α- and γ-tocopherols by 60% and 30%, respectively, promoted chlorophyll degradation and significantly enhanced VTE5 and VTE6 expression, and also that of HPPD and VTE4, altogether increasing total tocopherol accumulation. In conclusion, ABA increases promote the transcription of phytol recycling enzymes, which may contribute to vitamin E biosynthesis during fruit development in stone fruits like sweet cherries.

A gibberellin-assisted study of the transcriptional and hormonal changes occurring at floral transition in peach buds (Prunus persica L. Batsch).

BACKGROUND: Flower load in peach is an important determinant of final fruit quality and is subjected to cost-effective agronomical practices, such as the thinning, to finely balance the sink-source relationships within the tree and drive the optimal amount of assimilates to the fruits. Floral transition in peach buds occurs as a result of the integration of specific environmental signals, such as light and temperature, into the endogenous pathways that induce the meristem to pass from vegetative to reproductive growth. The cross talk and integration of the different players, such as the genes and the hormones, are still partially unknown. In the present research, transcriptomics and hormone profiling were applied on bud samples at different developmental stages. A gibberellin treatment was used as a tool to identify the different phases of floral transition and characterize the bud sensitivity to gibberellins in terms of inhibition of floral transition. RESULTS: Treatments with gibberellins showed different efficacies and pointed out a timeframe of maximum inhibition of floral transition in peach buds. Contextually, APETALA1 gene expression was shown to be a reliable marker of gibberellin efficacy in controlling this process. RNA-Seq transcriptomic analyses allowed to identify specific genes dealing with ROS, cell cycle, T6P, floral induction control and other processes, which are correlated with the bud sensitivity to gibberellins and possibly involved in bud development during its transition to the reproductive stage. Transcriptomic data integrated with the quantification of the main bioactive hormones in the bud allowed to identify the main hormonal regulators of floral transition in peach, with a pivotal role played by endogenous gibberellins and cytokinins. CONCLUSIONS: The peach bud undergoes different levels of receptivity to gibberellin inhibition. The stage with maximum responsiveness corresponded to a transcriptional and hormonal crossroad, involving both flowering inhibitors and inductors. Endogenous gibberellin levels increased only at the latest developmental stage, when floral transition was already partially achieved, and the bud was less sensitive to exogenous treatments. A physiological model summarizes the main findings and suggests new research ideas to improve our knowledge about floral transition in peach.

Biological outliers: essential elements to understand the causes and consequences of reductions in maximum photochemical efficiency of PSII in plants.

MAIN CONCLUSION: By studying Cistus albidus shrubs in their natural habitat, we show that biological outliers can help us to understand the causes and consequences of maximum photochemical efficiency decreases in plants, thus reinforcing the importance of integrating these often-neglected data into scientific practice. Outliers are individuals with exceptional traits that are often excluded of data analysis. However, this may result in very important mistakes not accurately capturing the true trajectory of the population, thereby limiting our understanding of a given biological process. Here, we studied the role of biological outliers in understanding the causes and consequences of maximum photochemical efficiency decreases in plants, using the semi-deciduous shrub C. albidus growing in a Mediterranean-type ecosystem. We assessed interindividual variability in winter, spring and summer maximum PSII photochemical efficiency in a population of C. albidus growing under Mediterranean conditions. A strong correlation was observed between maximum PSII photochemical efficiency (Fv/Fm ratio) and leaf water desiccation. While decreases in maximum PSII photochemical efficiency did not result in any damage at the organ level during winter, reductions in the Fv/Fm ratio were associated to leaf mortality during summer. However, all plants could recover after rainfalls, thus maximum PSII photochemical efficiency decreases did not result in an increased mortality at the organism level, despite extreme water deficit and temperatures exceeding 40ºC during the summer. We conclude that, once methodological outliers are excluded, not only biological outliers must not be excluded from data analysis, but focusing on them is crucial to understand the causes and consequences of maximum PSII photochemical efficiency decreases in plants.

Physiological mechanisms underlying extreme longevity in mountain pine trees.

Ancient trees are life history longevity winners that mostly persist in remote and environmentally harsh mountainous areas. Here, we performed a multifeature analysis in a protected mature mountain pine (Pinus uncinata) forest to identify the morphological and physiological traits that make these trees unique. We compared the physiology of meristematic and somatic tissues (apical buds and needles, respectively) from juvenile, mature young, mature old, and mature ancient trees under cold stress and nonstress conditions. We successfully identified key morphological features of extreme longevity at the organism level, as well as various growth, vigor, stress, and dormancy markers underlying extreme longevity in old and ancient trees. Results indicated that evolution has exerted selective pressure on specific physiological traits that make trees become longevity winners (<0.1% of the tree population were ancient trees, with an average trunk diameter >100 cm and an estimated age of 700 years). Traits entailing longevity not only included apical dominance loss, epicormic growth, and modular senescence, but also an extreme plasticity in both meristematic and somatic tissues (buds and needles, respectively), as shown by various physiological markers. In conclusion, ancient trees are oddities that not only possess a unique ecological value but also show divergent physiological behaviors selected during their evolution to allow them to cope with adversities and attain long life.

Experimental approaches in studying active biomolecules modulating fruit ripening: Melatonin as a case study.

Phytohormones are naturally occurring small organic molecules found at low concentrations in plants. They perform essential functions in growth and developmental processes, from organ initiation to senescence, including fruit ripening. These regulatory molecules are studied using different experimental approaches, such as performing exogenous applications, evaluating endogenous levels, and/or obtaining genetically modified lines. Here, we discuss the advantages and limitations of current experimental approaches used to study active biomolecules modulating fruit ripening, focusing on melatonin. Although melatonin has been implicated in fruit ripening in several model fruit crops, current knowledge is affected by the different experimental approaches used, which have given different and sometimes even contradictory results. The methods of application and the doses used have produced different results in studies based on exogenous applications, while different measurement methods and ways of expressing results explain most of the variability in studies using correlative analyses. Furthermore, studies on genetically modified crops have focused on tomato (Solanum lycopersicum L.) plants only. However, TILLING and CRISPR methodologies are becoming essential tools to complement the results from the experimental approaches described above. This will not only help the scientific community better understand the role of melatonin in modulating fruit ripening, but it will also help develop technological advances to improve fruit yield and quality in major crops. The combination of various experimental approaches will undoubtedly lead to a complete understanding of the function of melatonin in fruit ripening in the near future, so that this knowledge can be effectively transferred to the field.

Hormonal response to recurrent seasonal stress in coastal and mountain scabiouses growing in their natural habitat: linking ABA and jasmonates with photoprotection.

Plant species distribution across ecosystems is influenced by multiple environmental factors, and recurrent seasonal stress events can act as natural selection agents for specific plant traits and limit species distribution. For that, studies aiming at understanding how environmental constraints affect adaptive mechanisms of taxonomically closely related species are of great interest. We chose two Scabiosa species inhabiting contrasting environments: the coastal scabious S. atropurpurea, typically coping with hot-dry summers in a Mediterranean climate, and the mountain scabious S. columbaria facing cold winters in an oceanic climate. A set of functional traits was examined to assess plant performance in these congeneric species from contrasting natural habitats. Both S. atropurpurea and S. columbaria appeared to be perfectly adapted to their environment in terms of adjustments in stomatal closure, CO2 assimilation rate and water use efficiency over the seasons. However, an unexpected dry period during winter followed by the typical Mediterranean hot-dry summer forced S. atropurpurea plants to deploy a set of photoprotective responses during summer. Aside from reductions in leaf water content and Fv/Fm, photoprotective molecules (carotenoids, α-tocopherol and anthocyanins) per unit of chlorophyll increased, mostly as a consequence of a severe chlorophyll loss. The profiling of stress-related hormones (ABA, salicylic acid and jasmonates) revealed associations between ABA and the bioactive jasmonoyl-isoleucine with the underlying photoprotective response to recurrent seasonal stress in S. atropurpurea. We conclude that jasmonates may be used together with ABA as a functional trait that may, at least in part, help understand plant responses to recurrent seasonal stress in the current frame of global climate change.

Organ-specific responses during acclimation of mycorrhizal and non-mycorrhizal tomato plants to a mild water stress reveal differential local and systemic hormonal and nutritional adjustments.

MAIN CONCLUSION: Tomato plant acclimation to a mild water stress implied tissue-specific hormonal and nutrient adjustments, being the root one of the main modulators of this response. Phytohormones are key regulators of plant acclimation to water stress. However, it is not yet clear if these hormonal responses follow specific patterns depending on the plant tissue. In this study, we evaluated the organ-specific physiological and hormonal responses to a 14 day-long mild water stress in tomato plants (Solanum lycopersicum cv. Moneymaker) in the presence or absence of the arbuscular mycorrhizal fungus Rhizoglomus irregulare, a frequently used microorganism in agriculture. Several physiological, production, and nutritional parameters were evaluated throughout the experiments. Additionally, endogenous hormone levels in roots, leaves, and fruits at different developmental stages were quantified by ultrahigh-performance liquid chromatography coupled to tandem mass spectrometry (UHPLC-MS/MS). Water deficit drastically reduced shoot growth, while it did not affect fruit production. In contrast, fruit production was enhanced by mycorrhization regardless of the water treatment. The main tissue affected by water stress was the root system, where huge rearrangements in different nutrients and stress-related and growth hormones took place. Abscisic acid content increased in every tissue and fruit developmental stage, suggesting a systemic response to drought. On the other hand, jasmonate and cytokinin levels were generally reduced upon water stress, although this response was dependent on the tissue and the hormonal form. Finally, mycorrhization improved plant nutritional status content of certain macro and microelements, specially at the roots and ripe fruits, while it affected jasmonate response in the roots. Altogether, our results suggest a complex response to drought that consists in systemic and local combined hormonal and nutrient responses.

A phloem-localized Arabidopsis metacaspase (AtMC3) improves drought tolerance.

Increasing drought phenomena pose a serious threat to agricultural productivity. Although plants have multiple ways to respond to the complexity of drought stress, the underlying mechanisms of stress sensing and signaling remain unclear. The role of the vasculature, in particular the phloem, in facilitating inter-organ communication is critical and poorly understood. Combining genetic, proteomic and physiological approaches, we investigated the role of AtMC3, a phloem-specific member of the metacaspase family, in osmotic stress responses in Arabidopsis thaliana. Analyses of the proteome in plants with altered AtMC3 levels revealed differential abundance of proteins related to osmotic stress pointing into a role of the protein in water-stress-related responses. Overexpression of AtMC3 conferred drought tolerance by enhancing the differentiation of specific vascular tissues and maintaining higher levels of vascular-mediated transportation, while plants lacking the protein showed an impaired response to drought and inability to respond effectively to the hormone abscisic acid. Overall, our data highlight the importance of AtMC3 and vascular plasticity in fine-tuning early drought responses at the whole plant level without affecting growth or yield.

Photo-oxidative stress leading to oxylipin accumulation may trigger drought-induced leaf abscission in olive trees.

Olive trees shed their leaves under severe drought as a defence mechanism. Foliar drought-induced abscission is a programmed process occurring in a differentiated cell layer at the base of the petiole. Considering the antioxidant properties of vitamin E and its interplay with lipid peroxidation-derived jasmonates in abiotic stress responses, we hypothesized about their possible role in abscission signaling by forming a jasmonates-increasing basipetal gradient along the leaf up to the abscission zone. We exposed young olives trees to water withdrawal for 21 days, after which five leaf sections, from the apex to the petiole, were sampled on both attached and detached leaves of irrigated and water-stressed trees. We found that prolonged drought stress resulted in a sharp reduction in the photosystem II efficiency, chlorophyll and vitamin E contents in leaves, leading to photo-oxidative stress, reflected by the increase in lipid peroxidation. In addition, the content of chloroplast-derived oxylipins and phytohormones, such as jasmonoyl-isoleucine and salicylic acid, increased. At the same time, α-tocopherol decreased in the petiole of water-stressed attached leaves, suggesting a conditioning for the abscission process to occur. Although no differences were observed in petioles from attached and detached leaves, the dropped ones showed higher oxidative stress in the leaf blade. It is concluded that redox signaling through oxylipins accumulation may trigger leaf abscission in drought-stressed olive trees. Mechanical stress is, however, additionally needed to execute leaf abscission once the abscission zone is properly conditioned.

Water stress protection by the arbuscular mycorrhizal fungus Rhizoglomus irregulare involves physiological and hormonal responses in an organ-specific manner.

Arbuscular mycorrhizal fungi may alleviate water stress in plants. Although several protection mechanisms have already been described, little information is available on how these fungi influence the hormonal response to water stress at an organ-specific level. In this study, we evaluated the physiological and hormonal responses to water stress in above and below-ground tissues of the legume grass Trifolium repens colonized by the arbuscular mycorrhizal fungus Rhizoglomus irregulare. Plants were subjected to progressive water stress and recovery. Different leaf and root physiological parameters, as well as phytohormone levels, were quantified. Water-stressed mycorrhizal plants showed an improved water status and no photoinhibition compared to uncolonized individuals, while some stress markers like α-tocopherol and malondialdehyde content, an indicator of the extent of lipid peroxidation, transiently increased in roots, but not in leaves. Water stress protection exerted by mycorrhiza appeared to be related to a differential root-to-shoot redox signaling, probably mediated by jasmonates, and mycorrhization enhanced the production of the cytokinin trans-zeatin in both roots and leaves. Overall, our results suggest that mycorrhization affects physiological, redox and hormonal responses to water stress at an organ-specific level, which may eventually modulate the final protection of the host from water stress.

Global gene flow releases invasive plants from environmental constraints on genetic diversity.

When plants establish outside their native range, their ability to adapt to the new environment is influenced by both demography and dispersal. However, the relative importance of these two factors is poorly understood. To quantify the influence of demography and dispersal on patterns of genetic diversity underlying adaptation, we used data from a globally distributed demographic research network comprising 35 native and 18 nonnative populations of Plantago lanceolata Species-specific simulation experiments showed that dispersal would dilute demographic influences on genetic diversity at local scales. Populations in the native European range had strong spatial genetic structure associated with geographic distance and precipitation seasonality. In contrast, nonnative populations had weaker spatial genetic structure that was not associated with environmental gradients but with higher within-population genetic diversity. Our findings show that dispersal caused by repeated, long-distance, human-mediated introductions has allowed invasive plant populations to overcome environmental constraints on genetic diversity, even without strong demographic changes. The impact of invasive plants may, therefore, increase with repeated introductions, highlighting the need to constrain future introductions of species even if they already exist in an area.

Vitamin E protects from lipid peroxidation during winter stress in the seagrass Cymodocea nodosa.

MAIN CONCLUSION: Adjustments in the antenna size and α-tocopherol contents provide protection from sustained damage in leaves of a seagrass, while low vitamin E contents appear to be enough to protect rhizomes (which appear to be more cold tolerant than leaves). Despite low temperatures can adversely affect the proper growth and development of marine angiosperms, by, among other processes, increasing reactive oxygen species production and causing oxidative damage to lipid membranes, the role of vitamin E in seagrasses, such as Cymodocea nodosa has not been explored thus far. Here, we aimed to better understand the possible role of this chain-breaking (peroxyl radical-trapping) antioxidant in response to low temperatures, and most particularly in relation to the occurrence of photo-inhibition and lipid peroxidation. Low temperatures caused an important desiccation of leaves, but not of rhizomes, which were much more tolerant to cold stress than leaves. Cold stress during winter was associated with chlorophyll loss and transient photo-inhibition, as indicated by reversible reductions in the Fv/Fm ratio. Adjustments in pigment antenna size and vitamin E contents per unit of chlorophyll during winter may help protect the photosynthetic apparatus from sustained photo-inhibitory damage and lipid peroxidation events in leaves. Rhizomes also accumulated significant amounts of vitamin E, although to a much lesser extent than leaves, and kept protected from lipid peroxidation during winter, as indicated by malondialdehyde contents, a product from secondary lipid peroxidation. It is concluded that vitamin E can help protect both leaves and rhizomes from lipid peroxidation, although cold stress during winter can cause transient photo-inhibition of the photosynthetic apparatus, in C. nodosa.

Holoparasitic plant-host interactions and their impact on Mediterranean ecosystems.

Although photosynthesis is essential to sustain life on Earth, not all plants use sunlight to synthesize nutrients from carbon dioxide and water. Holoparasitic plants, which are important in agricultural and natural ecosystems, are dependent on other plants for nutrients. Phytohormones are crucial in holoparasitic plant-host interactions, from seed germination to senescence, not only because they act as growth and developmental regulators, but also because of their central role in the regulation of host photosynthesis and source-sink relations between the host and the holoparasitic plant. Here, we compile and discuss current knowledge on the impact and ecophysiology of holoparasitic plants (such as the broomrapes Orobanche sp. and Phelipanche sp.) that infest economically important dicotyledonous crops in Mediterranean agroecosystems (legumes [Fabaceae], sunflowers [Helianthus sp.], or tomato [Solanum lycopersicum] plants). We also highlight the role of holoparasitic plant-host interactions (such as those between Cytinus hypocistis and various shrubs of the genus Cistus) in shaping natural Mediterranean ecosystems. The roles of phytohormones in controlling plant-host interactions, abiotic factors in parasitism, and the biological significance of natural seed banks and how dormancy and germination are regulated, will all be discussed. Holoparasitic plants are unique organisms; improving our understanding of their interaction with hosts as study models will help us to better manage parasitic plants, both in agricultural and natural ecosystems.

Hormonal impact on photosynthesis and photoprotection in plants.

Photosynthesis is not only essential for plants, but it also sustains life on Earth. Phytohormones play crucial roles in developmental processes, from organ initiation to senescence, due to their role as growth and developmental regulators, as well as their central role in the regulation of photosynthesis. Furthermore, phytohormones play a major role in photoprotection of the photosynthetic apparatus under stress conditions. Here, in addition to discussing our current knowledge on the role of the phytohormones auxin, cytokinins, gibberellins, and strigolactones in promoting photosynthesis, we will also highlight the role of abscisic acid beyond stomatal closure in modulating photosynthesis and photoprotection under various stress conditions through crosstalk with ethylene, salicylates, jasmonates, and brassinosteroids. Furthermore, the role of phytohormones in controlling the production and scavenging of photosynthesis-derived reactive oxygen species, the duration and extent of photo-oxidative stress and redox signaling under stress conditions will be discussed in detail. Hormones have a significant impact on the regulation of photosynthetic processes in plants under both optimal and stress conditions, with hormonal interactions, complementation, and crosstalk being important in the spatiotemporal and integrative regulation of photosynthetic processes during organ development at the whole-plant level.

Phenotypic plasticity masks range-wide genetic differentiation for vegetative but not reproductive traits in a short-lived plant.

Genetic differentiation and phenotypic plasticity jointly shape intraspecific trait variation, but their roles differ among traits. In short-lived plants, reproductive traits may be more genetically determined due to their impact on fitness, whereas vegetative traits may show higher plasticity to buffer short-term perturbations. Combining a multi-treatment greenhouse experiment with observational field data throughout the range of a widespread short-lived herb, Plantago lanceolata, we (1) disentangled genetic and plastic responses of functional traits to a set of environmental drivers and (2) assessed how genetic differentiation and plasticity shape observational trait-environment relationships. Reproductive traits showed distinct genetic differentiation that largely determined observational patterns, but only when correcting traits for differences in biomass. Vegetative traits showed higher plasticity and opposite genetic and plastic responses, masking the genetic component underlying field-observed trait variation. Our study suggests that genetic differentiation may be inferred from observational data only for the traits most closely related to fitness.

Linking jasmonates with vitamin E accumulation in plants: a case study in the Mediterranean shrub Cistus albidus L.

MAIN CONCLUSION: Jasmonic acid positively modulates vitamin E accumulation, but the latter can also partly influence the capacity to accumulate the jasmonic acid precursor, 12-oxo-phytodienoic acid, in white-leaved rockrose (Cistus albidus L.) plants growing in their natural habitat. This study suggests a bidirectional link between chloroplastic antioxidants and lipid peroxidation-derived hormones in plants. While vitamin E is well known for its antioxidant properties being involved in plant responses to abiotic stress, jasmonates are generally related to biotic stress responses in plants. Studying them in non-model plants under natural conditions is crucial for the knowledge on their relationship, which will help us to better understand mechanisms and limits of stress tolerance to implement better conservation strategies in vulnerable ecosystems. We studied a typical Mediterranean shrub, white-leaved rockrose (Cistus albidus) under natural conditions during three winters and we analyzed both α and γ-tocopherol, and the three main jasmonates forms 12-oxo-phytodienoic acid (OPDA), jasmonic acid (JA), and jasmonoyl-isoleucine (JA-Ile). We found that JA contents positively correlated with vitamin E accumulation, most particularly with γ-tocopherol, the precursor of α-tocopherol (the most active vitamin E form). This finding was confirmed by exogenous application of methyl jasmonate (MeJA) in leaf discs under controlled conditions, which increased γ-tocopherol when applied at 0.1 mM MeJA and α-tocopherol at 1 mM MeJA. Furthermore, a complementary meta-analysis study with previously published reports revealed a positive correlation between JA and vitamin E, although this relationship turned to be strongly species specific. A strong negative correlation was observed, however, between total tocopherols and OPDA (a JA precursor located in chloroplasts). This antagonistic effect was observed between α-tocopherol and OPDA, but not between γ-tocopherol and OPDA. It is concluded that (i) variations in jasmonates and vitamin E due to yearly, inter-individual and sun orientation-driven variability are compatible with a partial regulation of vitamin E accumulation by jasmonates, (ii) vitamin E may also exert a role in the modulation of the biosynthesis of OPDA, with a much smaller effect, if any, on other jasmonates, and (iii) a trade-off in the accumulation of vitamin E and jasmonates might occur in the regulation of biotic and abiotic stress responses in plants.

Functional responses to climate change may increase invasive potential of Carpobrotus edulis.

PREMISE: Biological invasions and climate change are major threats to biodiversity. It is therefore important to anticipate how the climate changes projected for Southern Europe would affect the ecophysiological performance of the invasive South African plant, Carpobrotus edulis (ice plant or sour fig), and its capacity to undergo rapid adaptive evolution. METHODS: We manipulated the climate conditions in a field plot located on the island of Sálvora (northwest of the Iberian Peninsula) to establish a full factorial experiment with C. edulis plants transplanted from four native (southern African) and four invasive (northwestern Iberian Peninsula) populations. Throughout 14 months we measured growth and functional traits of this species under two temperatures (control vs. increased), and two rainfall levels (control vs. reduced). RESULTS: Temperature increased photochemical efficiency and relative growth rate of C. edulis. Rainfall modulated some of the effects of temperature on C and N isotopic composition, and pigment contents. Invasive populations showed lower root mass allocation and higher survival rates, as well as increased water use efficiency, lipid peroxidation, chlorophyll, and xanthophyll cycle pigment contents than native populations. CONCLUSIONS: The increased growth and physiological performances observed under our experimental conditions suggest that the expected climate changes would further promote the invasion of C. edulis. Differences between native and invasive genotypes in survival and functional traits revealed that populations have diverged during the process of invasion, what gives support to the invasiveness hypothesis. Our findings highlight the importance of analyzing intraspecific variability in functional responses to better predict how invasive species will respond to environmental changes.

English plantain deploys stress tolerance mechanisms at various organization levels across an altitudinal gradient in the Pyrenees.

High-mountain plants must withstand high solar irradiation and low temperatures during winter. Furthermore, climate change is increasing drought events, which pose an additional threat to plants. Here, we studied the stress tolerance mechanisms at various levels of biological organization in English plantain (Plantago lanceolata L.), focusing on photoprotective and antioxidant responses. The response of populations from three different altitudes in the Eastern Pyrenees (1030, 1380, and 1660 m. a.s.l.) was compared during both autumn and winter. Results showed that plants not only suffered from photoinhibition due to very low temperatures at the highest elevation during winter, but also from mild drought stress at the lowest altitude during autumn. Individuals growing at the highest elevation showed reductions in the maximum photochemical efficiency of PSII (Fv /Fm ratio), which might be caused by the lack of an increased induction of tolerance mechanisms at the highest elevation compared to the intermediate one. Although most leaves died at the highest elevation, plants could withstand stress at the organism level by generating new leaves once the stress ceased. Drought at the lowest elevation during autumn caused mild stress with small decreases in the Fv /Fm ratio, along with an increase in abscisic acid and jasmonic acid content. This study underlines the great capacity of English plantain to adapt to high elevation by activating not only photo- and antioxidant protection mechanisms and adjustments in stress-related phytohormones, but also by fully regenerating its aboveground biomass through renewed growth once the stress has ceased.

Geographic patterns of seed trait variation in an invasive species: how much can close populations differ?

Seeds play a major role in plant species persistence and expansion, and therefore they are essential when modeling species dynamics. However, homogeneity in seed traits is generally assumed, underestimating intraspecific trait variability across the geographic space, which might bias species success models. The aim of this study was to evaluate the existence and consequences of interpopulation variability in seed traits of the invasive species Carpobrotus edulis at different geographical scales. We measured seed production, morphology, vigour and longevity of nine populations of C. edulis along the Catalan coast (NE Spain) from three differentiated zones with a human presence gradient. Geographic distances between populations were contrasted against individual and multivariate trait distances to explore trait variation along the territory, evaluating the role of bioclimatic variables and human density of the different zones. The analysis revealed high interpopulation variability that was not explained by geographic distance, as regardless of the little distance between some populations (< 0.5 km), significant differences were found in several seed traits. Seed production, germination, and persistence traits showed the strongest spatial variability up to 6000% of percent trait variability between populations, leading to differentiated C. edulis soil seed bank dynamics at small distances, which may demand differentiated strategies for a cost-effective species management. Seed trait variability was influenced by human density but also bioclimatic conditions, suggesting a potential impact of increased anthropogenic pressure and climate shifts. Geographic interpopulation trait variation should be included in ecological models and will be important for assessing species responses to environmental heterogeneity and change.