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This study introduces a sustainable and environmentally friendly method for synthesizing selenium nanoparticles (SeNPs) by using Calluna vulgaris as a reducing agent. The process involves the addition of Na2SeO3 to a C. vulgaris aqueous solution, followed by reduction with ascorbic acid. UV-Vis spectroscopy confirmed SeNP formation, with a distinct absorption peak at 289 nm. Morphological analysis via Scanning Electron Microscopy (SEM) revealed spherical nanoparticles below 100 nm, as corroborated by Transmission Electron Microscopy (TEM) images displaying sizes ranging from 42.91 to 66.93 nm. Energy Dispersive Spectroscopy (EDS) confirmed the presence of selenium. Antibacterial assessments demonstrated the efficacy of C.vulgaris Selenium Nanoparticles (Cv-SeNPs) against gram-positive (Enterococcus faecalis, Staphylococcus aureus) and gram-negative bacteria (Escherichia coli). Cv-SeNPs exhibited notable antibacterial activity, particularly against E. Faecalis. In terms of antioxidant activities, Cv-SeNPs exhibited significant scavenging potential against DPPH and ABTS radicals, with low IC50 values of 24.72 and 16.87 µg/mL, respectively. The scavenging activities increased with concentration, reaching 86.6% for DPPH and 99.7% for ABTS at specific concentrations. The inclusion of ascorbic acid as a capping agent further augmented the free radical scavenging capabilities, indicating a synergistic relationship between selenium nanoparticles and capping agents. This research underscores the dual functionality of Cv-SeNPs as effective antibacterial agents and potent antioxidants. The green synthesis methodology utilizing C. vulgaris offers a sustainable approach for producing selenium nanoparticles with desirable characteristics, suggesting potential applications in medicine and industry. Further research on biomedical and industrial uses of Cv-SeNPs is needed.

Background: Predicting fire behaviour is an ongoing challenge in temperate peatlands and heathlands, where live fuels can form the dominant fuel load for wildfire spread, and where spatial heterogeneity in fuel moisture is important but not typically represented in fuel models. Aims: We examine the impact of fuel moisture variation on simulated fire behaviour across a temperate peatland/heathland landscape. Methods: We collected field measurements of fuel moisture content in Calluna vulgaris shrub from 36 sites across the North Yorkshire Moors, United Kingdom. We used these to define fuel moisture inputs within existing shrubland fuel models to simulate fire behaviour in BehavePlus. Key results: Simulated rates of spread varied with fuel moisture content; average mean variance of 23–80% from the landscape average rate of spread. The driest sites had simulated rates of spread up to 135% above the landscape average and the wettest sites up to 86% below average. Fuel model selection dramatically impacted simulated rates of spread by a factor of five. Conclusions: We need to constrain the role of live fuel moisture within temperate fuel models to develop accurate fire behaviour predictions. Implications: Capturing cross-landscape heterogeneity in fire behaviour is important for safe and effective land and wildfire management decision-making. Predicted fire behaviour varies significantly when cross-landscape fuel moisture differences are included in simulations. In temperate fuels, where live fuel moisture is not well represented in models, this may create dangerous situations where fire behaviour is underpredicted during the typical spring fire season, which has implications for fire management decision-making. [ABSTRACT FROM AUTHOR]

This study introduces a sustainable and environmentally friendly method for synthesizing selenium nanoparticles (SeNPs) by using Calluna vulgaris as a reducing agent. The process involves the addition of Na2SeO3 to a C. vulgaris aqueous solution, followed by reduction with ascorbic acid. UV-Vis spectroscopy confirmed SeNP formation, with a distinct absorption peak at 289 nm. Morphological analysis via Scanning Electron Microscopy (SEM) revealed spherical nanoparticles below 100 nm, as corroborated by Transmission Electron Microscopy (TEM) images displaying sizes ranging from 42.91 to 66.93 nm. Energy Dispersive Spectroscopy (EDS) confirmed the presence of selenium. Antibacterial assessments demonstrated the efficacy of C.vulgaris Selenium Nanoparticles (Cv-SeNPs) against gram-positive (Enterococcus faecalis, Staphylococcus aureus) and gram-negative bacteria (Escherichia coli). Cv-SeNPs exhibited notable antibacterial activity, particularly against E. Faecalis. In terms of antioxidant activities, Cv-SeNPs exhibited significant scavenging potential against DPPH and ABTS radicals, with low IC50 values of 24.72 and 16.87 µg/mL, respectively. The scavenging activities increased with concentration, reaching 86.6% for DPPH and 99.7% for ABTS at specific concentrations. The inclusion of ascorbic acid as a capping agent further augmented the free radical scavenging capabilities, indicating a synergistic relationship between selenium nanoparticles and capping agents. This research underscores the dual functionality of Cv-SeNPs as effective antibacterial agents and potent antioxidants. The green synthesis methodology utilizing C. vulgaris offers a sustainable approach for producing selenium nanoparticles with desirable characteristics, suggesting potential applications in medicine and industry. Further research on biomedical and industrial uses of Cv-SeNPs is needed. [ABSTRACT FROM AUTHOR]

For the frst time, AI algorithms based on drone imagery allow us to deduce quantitative statements on the reproductive behaviour of Calluna vulgaris individuals at the population level – especially with regard to their spatial distribution at several reproductive phases – which can be done individual-specifcally for a large number of individuals. The patterns thus extracted provide surprising insights into the variability of the fowering phenology of populations. For example, there is always an interplay within a population between profusely and weakly fowering individuals. However, their spatial confguration does not provide any recognizable patterns and its temporal appearance between the years does not remain constant. Even though a Calluna population reacts to changing environmental conditions with a median behaviour, i. e. a population fowers a lot or a little on average, no conclusions can be drawn from this as to the individual fowering behaviour. We are still unable to piece together the complex interplay of biotic and abiotic factors to draw a complete picture of the regulatory mechanisms in plant populations. In the future, the use of drones will allow us to generate further information and new knowledge about ecological relationships – in particular about the functioning of complex individual interactions in the realm of fowering plants and their effects on pollinating insects. At the same time, new AI-based evaluation methods are being developed that can map a variety of other plant characteristics and place them in the context of population dynamics so that – based on knowledge – the long-term protection and the development of natural habitats can be implemented. [ABSTRACT FROM AUTHOR]

Vegetation fuel dynamics in the UK's upland moorlands are important in determining landscape susceptibility to wildfire. Changes in fuel availability are influenced by phenology, land management activities or disturbances such as wildfires. Monitoring such changes is therefore essential to assess wildfire risks and impacts. This study used vegetation indices (VIs) derived from Sentinel-2 time series (2017–2023) and harmonic modelling to capture the phenology of key fuel properties and monitor post-fire vegetation recovery in four upland land cover types (acid grassland, heather, heather grassland, bog). We identified periods of high flammability, assessed the impact of wildfires on the spectral signal, and determined the time for spectral recovery as well as potential drivers of recovery times. Results showed the period of highest flammability from mid-February to early May in acid grassland, extending to early June in heather and heather grassland, and late June in bog. Summer fires caused more pronounced changes in fuel properties than spring fires, particularly in moisture-related VIs. Graminoid-dominated areas recovered rapidly (under a year), matching field observations, while dwarf shrub-dominated areas required up to three years, consistent with measurements of vegetation cover on burned areas but not with height. Spectral recovery times were primarily explained by land cover class, burn severity, season, and winter snow cover (R2 = 0.66). Field data highlighted pre-fire stand age's role in heather recovery and grasses' impact on spectral signals. This study improves understanding of fuel dynamics in upland moorlands through satellite monitoring, providing critical insights for more effective wildfire risk assessments and management strategies.

The ongoing loss of red‐listed coastal heathlands is a threat to biodiversity and cultural heritage legacies throughout the Atlantic coastal regions of Europe. It is possible to restore degraded and afforested heathlands, but restoration interventions are often labour‐intensive and costly, and the outcome of specific restoration actions are not well documented. We assess the efficiency of restoring coastal heathlands through natural succession (i.e. 'passive restoration') after removal of Sitka spruce Picea sitchensis (Bong.) Carr. plantations. The study was replicated on two neighbouring islands in a nature reserve in Western Norway. Low‐intensity free‐range sheep grazing was implemented as part of the reserve management plan. Furthermore, we tested the effect of leaving the clear‐felled woody material as chips on site, this being a cost‐efficient strategy on islands. Succession was monitored 1, 2, 4/5 and 8 years after clear‐felling, and revegetation of vascular plants and bryophytes was compared to target heathland vegetation. Surprisingly, we found different successional trajectories on the two islands. Species composition on one island approached target heathland vegetation during succession, but not on the other. Wood chips reduced species richness and slowed the restoration process, but these negative effects were only short‐term (<8 years). Differences in seed bank composition and soil conditions due to land use may explain the deviating successional trajectories on the two islands. We also found that management actions beyond clear‐felling and introducing sheep grazing are necessary due to the rapid seed regeneration of the Sitka spruce. [ABSTRACT FROM AUTHOR]

Atmospheric pollution containing soil‐nitrifying ammonium sulfate ((NH₄)₂SO₄) affects semi‐natural ecosystems worldwide. Long‐term additions of (NH₄)₂SO₄ to nitrogen (N)‐limited habitats, including heathlands, increase climate stress affecting recovery from wildfires. Although heathland vegetation largely depends on ericoid mycorrhizal fungi (ErM) to access soil N, we lack detailed understanding of how prolonged exposure to (NH₄)₂SO₄ may alter ErM community composition and host plants' reliance on fungal partners following wildfire and affect recovery. Simulation of atmospheric pollution ((NH₄)₂SO₄) occurred bi‐weekly for 5 years after a 2006 wildfire in a UK heathland. Ten years after treatments ceased, we measured vegetation structure, lichen and lichen photobiont composition, soil characteristics, ErM colonization, ErM diversity in roots and soil, and assessed ErM potential as novel recovery indicators. Heather height and density, and moss groundcover, were greater in N‐enriched plots. Lichen community indices showed significant treatment effects, but without differences in photobionts. Soil pH and Mg were significantly lower in treated plots while soil cation exchange capacity was significantly higher. There were no detectable differences in ErM composition and keystone ErM taxa between control and treated plots. Soil carbon stock measures were variable. Our results indicate atmospheric pollution following fire can have significant lingering effects above‐ and belowground. ErM diversity and root colonization were not assessed in the original N‐addition experiment; we advocate for their inclusion in future studies as an integral part of the recovery assessment toolkit. We show that mycorrhizal fungi diversity is a viable ecological tool and summarize key steps for ErM identification. [ABSTRACT FROM AUTHOR]

Background: Direct fuel moisture content measurements are critical for characterising spatio-temporal variations in fuel flammability and for informing fire danger assessments. However, among-sampler variability (systematic differences in measurements between samplers) likely contributes to fuel moisture measurement variability in most field campaigns. Aims: We assessed the magnitude of among-sampler variability in plot-scale Calluna vulgaris fuel moisture measurements. Methods: Seventeen individuals collected samples from six fuel layers hourly from 10:00 hours to 18:00 hours. We developed mixed effects models to estimate the among-sampler variability. Key results: Fuel moisture measurements were highly variable between individuals sampling within the same plot, fuel layer, and time of day. The importance of among-sampler variability in explaining total measured fuel moisture variance was fuel layer dependent. Among-sampler variability explained the greatest amount of measurement variation in litter (58%) and moss (45%) and was more important for live (19%) than dead (4%) Calluna. Conclusions: Both consideration of samplers within the experimental design and incorporation of sampler metadata during statistical analysis will improve understanding of spatio-temporal fuel moisture dynamics obtained from field-based studies. Implications: Accounting for among-sampler variability in fuel moisture campaigns opens opportunities to utilise sampling teams and citizen science research to examine fuel moisture dynamics over large spatio-temporal scales. Citizen science provides an important opportunity for the wildfire community, enabling data collection at broad spatio-temporal scales. Direct fuel moisture measurement campaigns can maximise these benefits by accounting for natural differences in measurements between individuals. We quantify the magnitude and variability of differences during an intensive fuel moisture measurement campaign. [ABSTRACT FROM AUTHOR]

Heathlands are unique cultural landscapes that once existed across vast stretches of northern Europe. Their deep-time persistence has formed an intrinsic part of economic and cultural practices. Such a complex interaction requires interdisciplinary approaches, including archaeology, across multiple regions to fully grasp all its aspects. The authors of this article review how research has been conducted in prehistoric heathlands across six nations in north-western Europe and outline the heaths' general characteristics. They discuss the major issues in that research, namely recurring narratives derived from history, an overall absence of consideration of the cultural aspects of heathlands, and a paucity of cross-regional initiatives. They suggest a series of theoretical and methodological approaches to improve this situation across expanded geographical and temporal scales. [ABSTRACT FROM AUTHOR]

Nitrogen (N) dynamic is one of the main controlling factors of responses to climate change in N‐limited terrestrial ecosystems, which rely on nutrient recycling and retention. In this study we investigate the N partitioning in ecosystem compartments of a grassland heath, and the impact of multiple climate change factors on long‐term N retention after 15N pulse labelling. The impacts of elevated carbon dioxide (eCO2), warming and drought and the treatments in combination on ecosystem N retention were investigated in a field scale manipulation experiment. A 6‐year time‐course was assessed by pulse‐labelling with the stable N isotope 15N and by sampling after 1 day, 1 year and 6 years. After 6 years we observed that the total ecosystem retained 42% of the amended 15N across treatments (recovery of the amended 15N in the pool). The fate of the applied 15N was mainly stabilization in soil, with 36% 15N recovery in soil, while the plant compartment and microbial biomass each retained only 1%–2% of the added 15N. This suggests a moderate retention of N, for all treatments, as compared to similar long‐term studies of forest ecosystems. A decreased ammonium and vegetation N pool combined with higher 15N retention in the soil under eCO2 treatments suggests that eCO2 promoted processes that immobilize N in soil, while warming counteracted this when combined with eCO2. Drought treatments contrastingly increased the vegetation N pool. We conclude that as the organic soil layer has the main capacity for N storage in a temperate heathland‐grassland, it is important for buffering nutrient availability and maintaining a resilient ecosystem. However, the full treatment combination of drought, warming and eCO2 did not differ in 15N recovery from the control, suggesting unchanged long‐term consequences of climate change on retention of pulse added N in this ecosystem. [ABSTRACT FROM AUTHOR]

The blanket bog ecosystems of the UK uplands fulfil important biodiversity and ecosystem services at multiple spatial scales. In response to perceived negative effects of burning to control heather growth on blanket bog, regulations on vegetation management now restrict this practice as part of the UK’s wider commitment to reducing carbon emissions. Instead, there has been a significant shift towards cutting as an alternative heather management tool, despite a notable lack of research on its effects on blanket bog. A key component of blanket bog ecosystems in the UK uplands is the moss layer, in which Sphagnum mosses play a particularly significant role. To investigate the immediate effects of heather cutting on the structure and extent of the moss layer, measurements of moss depth and cover at the end of the cutting season were taken from cut and uncut areas of two managed blanket bog sites in northern England. Measurements of moss depth were used to generate an index of moss microtopography. Cutting resulted in significant reductions of moss depth, heterogeneity of moss hummock microtopography, and extent of pleurocarpous moss, but there was no change in cover of Sphagnum or acrocarpous mosses. Further research is needed to understand the effects of cutting, particularly in the longer term, in terms of cutting height, type of machinery used, whether brash is removed, and repeat cut interval. This research will inform evidence-based management of blanket bog habitats.

Peatland ecosystems are of global conservation and environmental importance storing globally significant amounts of ancient carbon, regulating regional temperatures and hydrological regimes, and supporting unique biodiversity. Livestock grazing, land‐use change, drainage, nutrient and acid deposition, and wildfire threaten the composition and function of many peatlands including those in the uplands of the United Kingdom. Presently, little is known about either the short‐ or long‐term effects of wildfires within these systems in the UK. Our study aimed to evaluate how plant communities respond to wildfires across a range of vegetation communities, soil types, and burn severities. We evaluated wildfire burn severity using the ground‐based Composite Burn Index adapted for treeless peatlands. Using paired burned–unburned plots, we quantified differences in the abundance of plant families and functional groups, vegetation diversity, and community composition. Multivariate differences in composition between burned and unburned areas were used as an index of community resilience to fire. Plots in heathland communities with shallow organic soils burned at the highest severities and had the greatest reductions in plant diversity and richness. There were significant declines in plot‐scale species richness and diversity with increasing burn severity. Graminoids were resilient to fire whilst Ericaceae tended to increase with higher severity. Bryophyte composition was substantially altered—pleurocarpous species declined and acrocarpous species increased with greater burn severity. Community resilience was related to ground layer burn severity with higher burn severity driving greater changes in communities. Wildfire effects on temperate peatlands are a function of fire weather and site environmental and ecological characteristics. Management policy should ensure that the risk of severe wildfires is mitigated to protect ecosystem function and biodiversity. This will require system‐specific fire management prescriptions across the gradient of peatland soil and vegetation types. [ABSTRACT FROM AUTHOR]

Anthropogenic heathlands are semi‐natural ecosystems with a unique cultural and biodiversity value, considered worthy of preservation across most of the world. Their rate of loss, however, is alarming. Currently, we know little about the heathlands' actual span of resilience affordances and their association with abiotic and anthropogenic factors, including how much additional intervention they need to persist. Consequently, we are missing out on vital knowledge for conservation, management and the historical persistence of heathlands.This paper develops a method to assess the ecological resilience affordances of Atlantic postglacial heaths in the absence of human management. We use 12 existing cases of heathland succession to establish a four‐step resilience grade for each site, which we regress onto a series of explaining factors and use it in predicting heath resilience across postglacial Atlantic Northern Europe.We find that temperature, humidity, elevation and sandiness have a positive correlation with high heathland resilience. Our predictive mapping shows an uneven distribution of ecological heath resilience across Atlantic Northern Europe within an area of 1,000 × 1,200 km of 5 × 5 km resolution.Historic heathland distributions far exceed areas that afford high heath resilience, suggesting that heath distribution and persistence depend on both abiotic and anthropogenic factors.Policy implications: The map predicting the ecological resilience of Atlantic postglacial heaths can be used by managers working towards heath preservation and restoration to prioritize conservation efforts and to plan management practices across Atlantic Northern Europe. Together with the predictive model, it provides an important initial screening tool to assess heathland resilience in the absence of management as well as the impact of atmospheric nitrogen. The results are equally relevant for scholars who are interested in humans' role in increasing and decreasing ecosystem resilience. Our predictive method can be applied in other regions across the world by adding regionally specific variables. [ABSTRACT FROM AUTHOR]

Calluna vulgaris L. (heather) is a traditional medicinal plant with anti-inflammatory and calming activities that are determined by the notable amounts of phytochemicals. The evaluation of different populations of plants that accumulate great amounts of bioactive compounds are requisite for the quality determination of plant materials and medicinal and nutritional products. The assessment of natural resources from a phytogeographic point of view is relevant. Phytochemical analysis of heather samples was carried out using spectrophotometric methods and HPLC-PDA techniques, while antioxidant activity was determined using ABTS and FRAP assays. A significant diversification of phenolic and triterpenic compounds and antioxidant activity was determined in the heather samples collected in distinct habitats. Natural habitats, due to their characteristic chemical heterogeneity, lead to the diversity of indicators characterizing the quality of plant raw materials. Chlorogenic acid and hyperoside were found to be predominant among the phenolic compounds, while ursolic, oleanolic acids, and uvaol prevailed among the triterpenic compounds. Thus, these compounds can be suggested as phytochemical markers, characteristic of the heather raw material from central Europe. [ABSTRACT FROM AUTHOR]

Calluna vulgaris, belonging to the Ericaceae family, is an invasive plant that has widely spread from Europe all across Asia, North America, Australia and New Zealand. Being able to survive in rigid soil and environmental conditions, it is nowadays considered to be of high nature-conservation value. Known for its nutritional and medicinal properties, C. vulgaris stands out for its varied physiochemical composition, spotlighting a wide range of biological activity. Among the most important bioactive compounds identified in C. vulgaris, the phenolic components found in different parts of this herbaceous plant are the main source of its diverse pro-health properties (antioxidant, anti-inflammatory, antimicrobial, chemoprotective, etc.). Nonetheless, this plant exhibits an excellent nectariferous potential for social insects such as honeybees; therefore, comparing the bioactive compounds observed in the plant and in the final product of the beehive, namely honey, will help us understand and find new insights into the health benefits provided by the consumption of C. vulgaris-related products. Thus, the main interest of this work is to review the nutritional profile, chemical composition and biological activities of the C. vulgaris plant and its related honey in order to encourage the future exploration and use of this health-promoting plant in novel foods, pharmacological products and apitherapy. [ABSTRACT FROM AUTHOR]

Prescribed burning of vegetation is a land management approach that can be used to prevent or reduce the risk of wildfires yet burns can escape and be an ignition source for wildfires. In Scotland, muirburn is a form of prescribed burning used to encourage new growth of heather for grouse and livestock. Future regulation aims to mitigate potential negative impacts of muirburn on biodiversity, peat carbon stocks and wildfire risk. In this study, we assessed the spatial co-occurrence of muirburn and wildfires across Scotland. Evidence of muirburn was quantified using high-resolution aerial imagery from 2018 and wildfire occurrence was a composite of satellite and aerial detection covering 2015 to 2020. Fires were mapped on to dominant vegetation habitat types (acid grasslands, heather grasslands, heather heathlands, and peatbogs) and peat soil. Overall, 96 % of the total wildfire area occurred outside moorlands managed by muirburn, with wildfires covering 1.1 % of total moorland area. Heather grasslands, peatbogs and habitats on peat soils had a greater proportional area of wildfires than other moorland habitats and habitats on non-peat soils. In contrast, heather heathlands with non-peat soils had a greater proportional area of muirburn. We suggest the limited co-occurrence of muirburn and wildfires, may be due to fuel load reduction following muirburns, but may arise due to wildfire prevention strategies on estates and/or lower occurrence of recreational caused wildfires. In addition to muirburn regulation, our findings emphasise the need for policy, management, awareness and education outside of muirburn areas to reduce the risk of wildfires. • Mapped muirburn (prescribed burning) and wildfire area across Scottish moorlands • Overall, 96 % wildfire area occurs outside of moorland managed by muirburn • Between 2015 and 2020 wildfires covered 1.1 % of the moorland land area • Moorland vegetation and soil types differed in proneness to muirburn and wildfire [ABSTRACT FROM AUTHOR]

This paper presents new records and noteworthy data on the following taxa in SE Europe and adjacent regions: diatom alga Stauroneis neofossilis, parasitic fungus Anthracoidea arenariae, horsetail Equisetum hyemale, liverwort Harpanthus flotovianus, mosses Fissidens exilis and Rhizomnium punctatum, monocots Epipactis helleborine subsp. orbicularis, Himantoglossum calcaratum subsp. rumelicum and Schoenus nigricans and dicots Calluna vulgaris, Mahonia aquifolium and Willemetia stipitata subsp. albanica

Prescribed fire is one of the most widely-used management tools to recover encroached rangelands. Fire has been reported to cause changes in the soil physical and chemical properties. However, the legacy effects of former plant species on soil responses to fire remains unknown. The legacy effect of the former extant plant species on soil carbon (C) and nitrogen (N) fractionation distribution after prescribed burning in topsoil (0–5 cm and 5–10 cm) was investigated in Mediterranean shrublands in Montseny. We sampled soils under five vegetation patch types: Cytisus scoparius L., Calluna vulgaris L., Erica arborea L., Pteridium aquilinum L., and Cladonia biocrusts, pre- and post-burning. Multivariate analysis on soil C and N fractions showed that soils under the legume Cytisus and the biocrust were the most differentiated. Vegetation patch types tended to respond differently to burning, soils under Cytisus, Cladonia and Calluna showing the strongest response. Total C and N, and C and N in sand decreased after burning in the 0–5 cm soil layer. Conversely, C in silt, as well as N in clay and silt, increased with soil depth after burning. This study will be helpful for understanding ecological legacy effects and their possible consequences when planning prescribed burning. [ABSTRACT FROM AUTHOR]

The present paper gives new records of the following taxa in SE Europe and adjacent regions, together with significant data pertaining to them: the pathogenic fungus Arthrocladiella mougeotii; the liverwort Riccia frostii; the mosses Campylopus fragilis, C. introflexus and Tomentypnum nitens; the fern Allosorus persicus; the monocots Allium atropurpureum and Sisyrinchium montanum; and the dicots Calluna vulgaris and Santolina chamaecyparissus.

Abstract Recent developments in high‐resolution ecosystem mapping using unmanned aerial vehicles (UAV) open up promising perspectives for the monitoring of fine‐scale vegetation patterns and ecological functioning. In this study, we examine the potential of UAV imagery to track the structural composition and related phenological traits of the dwarf shrub Calluna vulgaris on a former military training area. On a European Natura 2000 heathland site, habitat management is shown to be evaluated on the basis of flowering dynamics and reproductive recovery as a proxy of functional changes that are generated after fire destruction. In particular, we utilize true color camera information and digital surface models to determine the spatiotemporal evolution of Calluna life‐cycle phases and flowering phenology 2 years after controlled burning. A stepwise methodological framework is presented that extracts Calluna pixels, spatially separates juvenile and mature life‐cycle phases, differentiates between generative and vegetative shoot extension and finally quantifies proportions of flowers, fruits and vegetative growth in two consecutive peak flowering periods. We show that Calluna life‐cycle phases can be spatially differentiated in pioneer, building and mature phases in UAV imagery. In the juvenile phase, regeneration from germination is low (

The blanket bog ecosystems of the UK uplands fulfil important biodiversity and ecosystem services at multiple spatial scales. In response to perceived negative effects of burning to control heather growth on blanket bog, regulations on vegetation management now restrict this practice as part of the UK's wider commitment to reducing carbon emissions. Instead, there has been a significant shift towards cutting as an alternative heather management tool, despite a notable lack of research on its effects on blanket bog. A key component of blanket bog ecosystems in the UK uplands is the moss layer, in which Sphagnum mosses play a particularly significant role. To investigate the immediate effects of heather cutting on the structure and extent of the moss layer, measurements of moss depth and cover at the end of the cutting season were taken from cut and uncut areas of two managed blanket bog sites in northern England. Measurements of moss depth were used to generate an index of moss microtopography. Cutting resulted in significant reductions of moss depth, heterogeneity of moss hummock microtopography, and extent of pleurocarpous moss, but there was no change in cover of Sphagnum or acrocarpous mosses. Further research is needed to understand the effects of cutting, particularly in the longer term, in terms of cutting height, type of machinery used, whether brash is removed, and repeat cut interval. This research will inform evidence-based management of blanket bog habitats. [ABSTRACT FROM AUTHOR]

Heather, Calluna vulgaris, is a key species of European dry heath and central determinant of its conservation status. The established Calluna life cycle concept describes four phases—pioneer, building, mature, and degeneration—distinguishable by growth and vitality characteristics of undisturbed plants grown from seeds. However, little is known about the life cycle and ageing of plants subjected to severe disturbance, although measures to this effect (burning, mowing) are common in heathland management. We studied the vitality of over 400 heather plants by examining multiple morphological (plant height, long shoot and inflorescence lengths, flowering activity), anatomical (growth rings) and environmental (management, nitrogen deposition, climate) attributes. We found Calluna vitality to be mainly determined by the aboveground stem age, and that severe disturbances promote vigorous vegetative regeneration. Ageing-related shifts in the habit and vitality of plants resprouting from stem-base buds is similar to that of seed-based plants, but the former revealed higher vitality when young, at the cost of a shorter life span. In contrast, plants originating from decumbent stems resemble building-stage plants but apparently lack the capacity to re-enter a cycle including stages other than degeneration-type. As a consequence, we supplemented the established heather life cycle concept with a post-disturbance regeneration cycle of plants derived from resprouting. We conclude that management of dry lowland heathlands should include rotational small-scale severe disturbance to support both seed germination and seedling establishment as well as vegetative regeneration chiefly of young heather plants capable of resprouting from buds near rootstock. [ABSTRACT FROM AUTHOR]

The Calluna vulgaris honey produced in Portugal, concerning its phenolic compounds and abscisic acids profiles, as well as its antioxidant activity and the protective effect against oxidative damage in human erythrocytes were herein performed for the first time. The phenolic and abscisic acid profiles were tentatively identified by LC-MS/MS (17 compounds). The total content of phenolics and abscisic acids was 15,446.4 μg/g of honey extract, with catechin derivatives and abscisic acids being major constituents. The highest scavenging capacity was found against reactive nitrogen species. Additionally, the honey extract prevented ROO'-induced oxidative damage in erythrocytes collected from human blood, by inhibiting hemolysis, lipid peroxidation and hemoglobin oxidation. In conclusion, C. vulgaris honey contains high content of catechin derivatives and abscisic acids that may be responsible for its biological activity, characterized by a strong antioxidant capacity, which adds up to the nutritional value of this delicacy. [ABSTRACT FROM AUTHOR]

Winter temperatures are projected to increase in Central Europe. Subsequently, snow cover will decrease, leading to increased soil temperature variability, with potentially different consequences for soil frost depending on e.g. altitude. Here, we experimentally evaluated the effects of increased winter soil temperature variability on the root associated mycobiome of two plant species (Calluna vulgaris and Holcus lanatus) at two sites in Germany; a colder and wetter upland site with high snow accumulation and a warmer and drier lowland site, with low snow accumulation. Mesocosm monocultures were set-up in spring 2010 at both sites (with soil and plants originating from the lowland site). In the following winter, an experimental warming pulse treatment was initiated by overhead infrared heaters and warming wires at the soil surface for half of the mesocosms at both sites. At the lowland site, the warming treatment resulted in a reduced number of days with soil frost as well as increased the average daily temperature amplitude. Contrary, the treatment caused no changes in these parameters at the upland site, which was in general a much more frost affected site. Soil and plant roots were sampled before and after the following growing season (spring and autumn 2011). High-throughput sequencing was used for profiling of the root-associated fungal (ITS marker) community (mycobiome). Site was found to have a profound effect on the composition of the mycobiome, which at the upland site was dominated by fast growing saprotrophs (Mortierellomycota), and at the lowland site by plant species-specific symbionts (e.g. Rhizoscyphus ericae and Microdochium bolleyi for C. vulgaris and H. lanatus respectively). The transplantation to the colder upland site and the temperature treatment at the warmer lowland site had comparable consequences for the mycobiome, implying that winter climate change resulting in higher temperature variability has large consequences for mycobiome structures regardless of absolute temperature of a given site. [ABSTRACT FROM AUTHOR]

Calluna vulgaris L. (heather) is a traditional medicinal plant with anti-inflammatory and calming activities that are determined by the notable amounts of phytochemicals. The evaluation of different populations of plants that accumulate great amounts of bioactive compounds are requisite for the quality determination of plant materials and medicinal and nutritional products. The assessment of natural resources from a phytogeographic point of view is relevant. Phytochemical analysis of heather samples was carried out using spectrophotometric methods and HPLC-PDA techniques, while antioxidant activity was determined using ABTS and FRAP assays. A significant diversification of phenolic and triterpenic compounds and antioxidant activity was determined in the heather samples collected in distinct habitats. Natural habitats, due to their characteristic chemical heterogeneity, lead to the diversity of indicators characterizing the quality of plant raw materials. Chlorogenic acid and hyperoside were found to be predominant among the phenolic compounds, while ursolic, oleanolic acids, and uvaol prevailed among the triterpenic compounds. Thus, these compounds can be suggested as phytochemical markers, characteristic of the heather raw material from central Europe.

Calluna vulgaris, belonging to the Ericaceae family, is an invasive plant that has widely spread from Europe all across Asia, North America, Australia and New Zealand. Being able to survive in rigid soil and environmental conditions, it is nowadays considered to be of high nature-conservation value. Known for its nutritional and medicinal properties, C. vulgaris stands out for its varied physiochemical composition, spotlighting a wide range of biological activity. Among the most important bioactive compounds identified in C. vulgaris, the phenolic components found in different parts of this herbaceous plant are the main source of its diverse pro-health properties (antioxidant, anti-inflammatory, antimicrobial, chemoprotective, etc.). Nonetheless, this plant exhibits an excellent nectariferous potential for social insects such as honeybees; therefore, comparing the bioactive compounds observed in the plant and in the final product of the beehive, namely honey, will help us understand and find new insights into the health benefits provided by the consumption of C. vulgaris-related products. Thus, the main interest of this work is to review the nutritional profile, chemical composition and biological activities of the C. vulgaris plant and its related honey in order to encourage the future exploration and use of this health-promoting plant in novel foods, pharmacological products and apitherapy.

The future status of peatlands as carbon stores/sinks is uncertain given current and predicted environmental change. Several factors can affect the magnitude of the peatland carbon sink including disturbances such as wildfire. There is at present little evidence of how wildfire affects the emission of carbon dioxide (CO2) and methane (CH4) via perturbation to aerobic and anaerobic respiration. The greatest effects, which are likely to vary according to wildfire severity, would be expected in the immediate post-fire stages when little recovery has taken place. Here, we investigate five UK peatland wildfires (2011–2012) in the immediate post-wildfire period measuring CO2 and CH4 fluxes using static chambers. Fire severity was described using a modified form of the composite burn index. A hierarchical partitioning approach indicated time since fire was the most strongly associated variable that fluxes of both CO2, and CH4 followed by soil temperature for CO2 and fire severity for CH4. Using a liner mixed modelling approach to account for repeated measures; fire severity was a significant term for CH4 and borderline significant for CO2. Mean fluxes of CH4 were consistently lower on burnt sites. In contrast, data from a fire in the north of Scotland appeared to show the opposite relationship for CH4 with higher fluxes on the burnt sites. These results suggest that wildfire can affect gaseous carbon fluxes, but the responses can be variable in both space and time and that disruption to anaerobic processes may be site and/or fire dependent. [ABSTRACT FROM AUTHOR]

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Questions: To improve our knowledge on how environmental conditions determine the development of high‐value Calluna vulgaris heathland habitats, we studied the floristic and structural characteristics of heathland plant communities across North Germany and how they are influenced by edaphic, climatic and management factors. We ask how heathland development is related to these factors and what are the implications for conservation management and restoration. Location: North German Plain. Methods: We collected 350 relevés in 18 dry Calluna heathland areas. Plant communities were classified using Isopam, and Redundancy Analysis (RDA) determined effects of environmental conditions. Potential pathways of development and the nature conservation status of the communities were identified on a multifactorial basis. Results: We found nine floristically and structurally distinct heathland plant communities. Heathland vegetation showed distinct patterns along Calluna age development stages and environmental conditions. Soil conditions and related effects of long‐term site history and recent management turned out to be the predominant factors influencing species composition and diversity, resulting in three potential heathland succession pathways. Mosaic‐like communities with particularly high taxonomic diversity and conservation value occurred on early‐successional inland dunes or as regeneration stage growing on nutrient‐poor sandy soils without humus accumulation. Conclusions: The study reveals fundamental differences between historically farmed heathland in the oceanic Northwest and former military training areas mainly in northeastern Germany with consequences for restoration ecology. Present nature conservation criteria turned out to be insufficient in predicting habitat quality, as lichens are frequently disregarded. Our findings highlight the need for intense soil disturbance to maintain early‐stage soil conditions and a diverse Calluna growth‐phase composition, as these factors essentially determine species richness in lowland heaths. [ABSTRACT FROM AUTHOR]

Background and Aims Plant secondary metabolites play critical roles in plant stress tolerance and adaptation, and are known to be influenced by the environment and climate changes, yet the impacts and interactions of multiple climate change components are poorly understood, particularly under natural conditions. Methods Accumulation of phenolics and emissions of volatile organic compounds (VOCs) were assessed on heather, Calluna vulgaris , an abundant evergreen dwarf shrub in European heathlands, after 6 years of exposure to elevated CO2, summer drought and nighttime warming. Key Results Drought alone had the strongest effects on phenolic concentrations and compositions, with moderate effects of elevated CO2 and temperature. Elevated CO2 exerted the greatest impact on VOC emissions, mainly by increasing monoterpene emissions. The response magnitudes varied among plant tissue types and chemical constituents, and across time. With respect to interactive effects of the studied climate change components, the interaction between drought and elevated CO2 was most apparent. Drought mainly reduced phenolic accumulation and VOC emissions, while elevated CO2 mitigated such effects. Conclusions In natural ecosystems, co-occurring climate factors can exert complex impacts on plant secondary metabolite profiles, which may in turn alter ecosystem processes. [ABSTRACT FROM AUTHOR]

Prunus serotina is one of the worst invasive alien plants throughout Europe, able to rapidly replace autochthonous vegetation and form monospecific stands both in open lands and forested areas. Invasion by shrubs and trees due to abandonment of traditional management practices, is one of the major threats to the conservation of "European dry heaths", a priority habitat included in the Council Directive 92/43/EEC. The aim of this study was to assess the effects of P. serotina invasion on heathland vegetation in terms of species composition and cover, with special regards to the dominant Calluna vulgaris and Molinia arundinacea, in an area of North-Western Italy subjected to massive invasion by this alien species. We collected data on the plant cover along an invasion gradient in 10 linear transects running from P. serotina invaded stands towards non invaded ones. P. serotina differently affected the heathland vegetation. In particular, the cover of C. vulgaris and other typical heathland species such as Cytisus scoparius sharply decreased at P. serotina cover increasing. On the contrary, positive or no effect of P. serotina cover was found on M. arundinacea. Our results showed that shaded conditions created by the alien tree disfavored light demanding C. vulgaris and C. scoparius, enabling fast grower M. arundinacea to outcompete and replace them. This study emphasizes the urgency to develop management strategies to effectively control P. serotina spread in the area and to restore typical heathland species. [ABSTRACT FROM AUTHOR]

Recent developments in high‐resolution ecosystem mapping using unmanned aerial vehicles (UAV) open up promising perspectives for the monitoring of fine‐scale vegetation patterns and ecological functioning. In this study, we examine the potential of UAV imagery to track the structural composition and related phenological traits of the dwarf shrub Calluna vulgaris on a former military training area. On a European Natura 2000 heathland site, habitat management is shown to be evaluated on the basis of flowering dynamics and reproductive recovery as a proxy of functional changes that are generated after fire destruction. In particular, we utilize true color camera information and digital surface models to determine the spatiotemporal evolution of Calluna life‐cycle phases and flowering phenology 2 years after controlled burning. A stepwise methodological framework is presented that extracts Calluna pixels, spatially separates juvenile and mature life‐cycle phases, differentiates between generative and vegetative shoot extension and finally quantifies proportions of flowers, fruits and vegetative growth in two consecutive peak flowering periods. We show that Calluna life‐cycle phases can be spatially differentiated in pioneer, building and mature phases in UAV imagery. In the juvenile phase, regeneration from germination is low (<10%) and sprouting from the stem base dominates recovery patterns after fire management. The study further reveals that the distribution of flowering and fruiting during peak flowering time is spatially heterogeneous with high variations between years. In the fire unaffected mature phase, proportions of flowers and fruits are reduced indicating a decrease in reproductive capacity of unmanaged Calluna stands. A phenological phase shift is detected from flowering to fruiting that is related to an increase of Calluna growth height. The study shows that the functional patterns of growth and reproduction can be mapped for heathland ecosystems using simple true color drone cameras. [ABSTRACT FROM AUTHOR]

Gastrointestinal nematode (GIN) infections impact small ruminant health, welfare, and production across farming systems. Rising anthelmintic resistance and regulation of synthetic drug use in organic farming is driving research and development of sustainable alternatives for GIN control. One alternative is the feeding of plants that contain secondary metabolites (PSMs) e.g., proanthocyanidins (PA, syn. condensed tannins) that have shown anthelmintic potential. However, PSMs can potentially impair performance, arising from reduced palatability and thus intake, digestibility or even toxicity effects. In this study, we tested the trade-off between the antiparasitic and anti-nutritional effects of heather consumption by lambs. The impact of additional feeding of a nematophagous fungus (Duddingtonia flagrans) on larval development was also explored. Lambs infected with Teladorsagia circumcincta or uninfected controls, were offered ad libitum heather, or a control chopped hay for 22 days during the infection patent period. Eight days into the patent period, parasitised lambs were supplemented (or remained unsupplemented) with D. flagrans for a 5-day period. Performance and infection metrics were recorded, and polyphenol levels in the heather and control hay were measured to investigate their association with activity. The lambs consumed heather at approximately 20% of their dry matter intake, which was sufficient to exhibit significant anthelmintic effects via a reduction in total egg output (P = 0.007), compared to hay-fed lambs; the magnitude of the reduction over time in heather fed lambs was almost 10-fold compared to control lambs. Negative effects on production were shown, as heather-fed lambs weighed 6% less than hay-fed lambs (P < 0.001), even though dry matter intake (DMI) of heather increased over time. D. flagrans supplementation lowered larval recovery in the faeces of infected lambs by 31.8% (P = 0.003), although no interactions between feeding heather and D. flagrans were observed (P = 0.337). There was no significant correlation between PA, or other polyphenol subgroups in the diet and egg output, which suggests that any association between heather feeding and anthelmintic effect is not simply and directly attributable to the measured polyphenols. The level of heather intake in this study showed no antagonistic effects on D. flagrans , demonstrating the methods can be used in combination, but provide no additive effect on overall anthelmintic efficacies. In conclusion, heather feeding can assist to reduce egg outputs in infected sheep, but at 20% of DMI negative effects on lamb performance can be expected which may outweigh any antiparasitic benefits. • Heather intake at 20% of DMI reduced gastrointestinal nematode daily egg output. • Adult worm counts were not affected by heather consumption in infected lambs. • Lambs on the heather diet suffered performance costs shown by lower bodyweight. • Supplementation with Duddingtonia flagrans reduced larval recovery in faeces. [ABSTRACT FROM AUTHOR]

The Calluna vulgaris honey produced in Portugal, concerning its phenolic compounds and abscisic acids profiles, as well as its antioxidant activity and the protective effect against oxidative damage in human erythrocytes were herein performed for the first time. The phenolic and abscisic acid profiles were tentatively identified by LC-MS/MS (17 compounds). The total content of phenolics and abscisic acids was 15,446.4 µg/g of honey extract, with catechin derivatives and abscisic acids being major constituents. The highest scavenging capacity was found against reactive nitrogen species. Additionally, the honey extract prevented ROO•-induced oxidative damage in erythrocytes collected from human blood, by inhibiting hemolysis, lipid peroxidation and hemoglobin oxidation. In conclusion, C. vulgaris honey contains high content of catechin derivatives and abscisic acids that may be responsible for its biological activity, characterized by a strong antioxidant capacity, which adds up to the nutritional value of this delicacy.

На основе литературных данных и собственных наблюдений разработана шкала оценки декоративности Calluna vulgaris (L.) Hull и его культиваров. Она содержит девять критериев, каждому из которых присвоен коэффициент значимости. Шкала оценки декоративности разделена на такие категории, как культивары с очень низкой (24-38 баллов), низкой (39-52), средней (53-66), повышенной (67-80) и высокой декоративностью (81-95). По разработанной шкале проведена оценка декоративности C. vulgaris и 10 культиваров, которые растут в коллекции НБС имени Н.Н. Гришко НАНУ в г. Киев, Украина. Выявлены культивары с высокой и повышенной декоративностью.

Copyright of Botanica Serbica is the property of University of Belgrade, Institute of Botany & Botanical Garden Jevremovac and its content may not be copied or emailed to multiple sites or posted to a listserv without the copyright holder's express written permission. However, users may print, download, or email articles for individual use. This abstract may be abridged. No warranty is given about the accuracy of the copy. Users should refer to the original published version of the material for the full abstract. (Copyright applies to all Abstracts.)

Peatlands represent a globally important carbon stock. Peat soil carbon stock assessments rely on measurements of carbon concentration, bulk density and soil depth. However, peat surface levels - and thus soil depths - change ("bog breathing") largely as a response to peat shrinkage and expansion in relation to water table changes. This study assessed the magnitude of and potential for this mechanism to influence carbon stock calculations under field and laboratory conditions. In the UK, most peatlands are upland blanket bogs, of which a large proportion are managed as 'grouse moors'. This currently involves rotational burning, although alternative management is increasingly considered. Peat depth and water tables were investigated on three grouse moors on blanket bog in northern England, comparing changes under different heather management interventions (burning, mowing and uncut). One site also allowed investigation of changes in relation to slope and under the three major bog vegetation types (ling heather, cottongrass and Sphagnum moss), which were compared to observed changes in peat cores under controlled laboratory conditions simulating periods of drought and rewetting. Changes in depth and bulk density were recorded and the potential implications for carbon stock estimates were calculated. Results highlight site specific relationships as potential habitat condition indicators and demonstrate that previously reported surface peat carbon losses resulting from enhanced decomposition under rising temperatures might also be explained by apparent changes due to peat shrinkage and expansion. It is recommended to accurately record bulk density and total peat depth measurements as part of peat carbon stock assessments. [ABSTRACT FROM AUTHOR]

Extreme climatic events are among the drivers of recent declines in plant biomass and productivity observed across Arctic ecosystems, known as "Arctic browning." These events can cause landscape‐scale vegetation damage and so are likely to have major impacts on ecosystem CO2 balance. However, there is little understanding of the impacts on CO2 fluxes, especially across the growing season. Furthermore, while widespread shoot mortality is commonly observed with browning events, recent observations show that shoot stress responses are also common, and manifest as high levels of persistent anthocyanin pigmentation. Whether or how this response impacts ecosystem CO2 fluxes is not known. To address these research needs, a growing season assessment of browning impacts following frost drought and extreme winter warming (both extreme climatic events) on the key ecosystem CO2 fluxes Net Ecosystem Exchange (NEE), Gross Primary Productivity (GPP), ecosystem respiration (Reco) and soil respiration (Rsoil) was carried out in widespread sub‐Arctic dwarf shrub heathland, incorporating both mortality and stress responses. Browning (mortality and stress responses combined) caused considerable site‐level reductions in GPP and NEE (of up to 44%), with greatest impacts occurring at early and late season. Furthermore, impacts on CO2 fluxes associated with stress often equalled or exceeded those resulting from vegetation mortality. This demonstrates that extreme events can have major impacts on ecosystem CO2 balance, considerably reducing the carbon sink capacity of the ecosystem, even where vegetation is not killed. Structural Equation Modelling and additional measurements, including decomposition rates and leaf respiration, provided further insight into mechanisms underlying impacts of mortality and stress on CO2 fluxes. The scale of reductions in ecosystem CO2 uptake highlights the need for a process‐based understanding of Arctic browning in order to predict how vegetation and CO2 balance will respond to continuing climate change. Extreme climatic events are among the drivers of vegetation damage and decline observed across Arctic ecosystems in recent years; termed "Arctic browning." Although these events can cause landscape‐scale vegetation damage, their impacts on ecosystem CO2 balance are little understood. Here, it is demonstrated that these events can have major impacts on CO2 balance, considerably reducing the carbon sink capacity of the ecosystem across the growing season. These impacts can be similar when associated with shoot mortality, or with shoot stress responses. [ABSTRACT FROM AUTHOR]

This study evaluated the interactive effects of combined phosphorus (P) and potassium (K) additions on canopy nitrogen (N) retention (CNR) and subsequent canopy leaching at a long-term N manipulation site on Whim bog in south Scotland. Ambient deposition is 8 kg N ha−1 year−1 and an additional 8, 24, and 56 kg N ha−1 year−1 of either ammonium (NH4+) or nitrate (NO3−) with or without P and K has been applied over 11 years. Throughfall N deposition below Calluna vulgaris and foliar N and P concentrations were assessed. Results showed that 60% for low dose and 53% for high dose of NO3− contrasting with 80% for low dose and 38% for high dose of NH4+ onto Calluna was retained by Calluna canopy. The CNR was enhanced by P and K addition in which 84% of NO3− and 83% of NH4+ for high dose were retained. CNR for NO3− increased the canopy leaching of dissolved organic N (DON) and associated organic anions. NH4+ retention increased canopy leaching of magnesium and calcium through ion exchange. Even over 11-years N exposure without P and K, foliage N:P ratio of Calluna did not increase, suggesting that N exposure did not lead to N saturation of Calluna at Whim bog. Our study concluded that increases in P and K availability enhance CNR of Calluna but accelerate the associated canopy leaching of DON and base cations, depending on foliar N status. [ABSTRACT FROM AUTHOR]

Understanding the climatic and biological factors that regulate soil carbon dioxide (CO2) efflux is crucial in peatlands because they contain a large proportion of terrestrial carbon (C). We predicted that rainfall reduction would increase soil CO2 efflux, and that cessation of below-ground allocation of recent plant assimilate would reduce soil CO2 efflux. These predictions were tested in the field using rainfall shelters that allowed a maximum of 40 % of rainfall onto 2 × 2 m plots by diverting rainwater from the shelter roofs with guttering, and by girdling stems of the dominant plant, Calluna vulgaris, for two years. We also used 13CO2-pulse labelling of intact monoliths at ambient CO2 concentrations to trace recent assimilate from plant shoots to roots, bulk soil, leachate, dissolved organic carbon (DOC) and soil CO2 efflux . Soil CO2 efflux in the sheltered plots increased in Year 1 but not in Year 2, and we found a positive relationship between soil CO2 efflux and water table depth. Our data indicate that lowering the water table below a critical threshold (15–20 cm) affects soil CO2 efflux. Girdling of C. vulgaris shoots resulted in no measurable reduction in soil CO2 efflux, while only ~3 % of 13C fixed by shoots was recovered in soil CO2 efflux and DOC in the 20 days after labelling. Our findings show that below-ground allocation of recent assimilate from C. vulgaris plants > 6 years old has little impact on soil CO2 efflux.