Synchrotron elemental imaging reveals zinc distribution in the hyperaccumulator Sedum plumbizincicola (Crassulaceae)

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Sedum plumbizincicola is a zinc-cadmium (Zn-Cd) hyperaccumulator native to China with high potential for use in phytoremediation of contaminated soils in temperate climate. This study aimed to determine the Zn accumulation and distribution in S. plumbizincicola tissues grown on soils co-contaminated with Cd, Pb and Zn. The efficiency of Zn accumulation was assessed in a monoculture and in co-cropping systems with Noccaea caerulescens. The samples were analysed by inductively coupled plasma–atomic emission spectrometry and synchrotron micro-X-ray fluorescence elemental imaging. Sedum plumbizincicola grown in monoculture showed significantly higher foliar concentrations of Zn than the plants grown with N. caerulescens, with the leaf tips, petioles and nodes being the main sites of Zn localisation in the aerial parts. The highest Zn concentrations were observed in the epidermis and vascular system, of both leaves and stems, with distribution pattern differing in young and in mature leaves. This study highlights the Zn localisation patterns in S. plumbizincicola to improve our understanding of the mechanisms of hyperaccumulation. Growing in monoculture, S. plumbizincicola is an effective candidate for Zn phytoremediation in contaminated soils, with less promising results when co-cropped with the Ganges ecotype of N. caerulescens.
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Synchrotron elemental imaging reveals zinc distribution in the hyperaccumulator Sedum plumbizincicola (Crassulaceae) | Authorea try { document.documentElement.classList.add('js'); } catch (e) { } var _gaq = _gaq || []; _gaq.push(['_setAccount', 'G-8VDV14Y67G']); _gaq.push(['_trackPageview']); (function() { var ga = document.createElement('script'); ga.type = 'text/javascript'; ga.async = true; ga.src = ('https:' == document.location.protocol ? 'https://ssl' : 'http://www') + '.google-analytics.com/ga.js'; var s = document.getElementsByTagName('script')[0]; s.parentNode.insertBefore(ga, s); })(); Skip to main content Preprints Collections Wiley Open Research IET Open Research Ecological Society of Japan All Collections About About Authorea FAQs Contact Us Quick Search anywhere Search for preprint articles, keywords, etc. Search Search ADVANCED SEARCH SCROLL Ecological Research This is a preprint and has not been peer reviewed. Data may be preliminary. 4 August 2025 V1 Latest version Share on Synchrotron elemental imaging reveals zinc distribution in the hyperaccumulator Sedum plumbizincicola (Crassulaceae) Authors : Julien Jacquet , Ksenija Jakovljević 0000-0002-1457-6807 , Dennis Brückner 0000-0003-1714-5452 , Catherine SIRGUEY 0000-0002-3181-9014 , and Antony van der Ent 0000-0003-0922-5065 [email protected] Authors Info & Affiliations https://doi.org/10.22541/au.175429444.47523987/v1 259 views 170 downloads Contents Abstract Information & Authors Metrics & Citations View Options References Figures Tables Media Share Abstract Sedum plumbizincicola is a zinc-cadmium (Zn-Cd) hyperaccumulator native to China with high potential for use in phytoremediation of contaminated soils in temperate climate. This study aimed to determine the Zn accumulation and distribution in S. plumbizincicola tissues grown on soils co-contaminated with Cd, Pb and Zn. The efficiency of Zn accumulation was assessed in a monoculture and in co-cropping systems with Noccaea caerulescens. The samples were analysed by inductively coupled plasma–atomic emission spectrometry and synchrotron micro-X-ray fluorescence elemental imaging. Sedum plumbizincicola grown in monoculture showed significantly higher foliar concentrations of Zn than the plants grown with N. caerulescens, with the leaf tips, petioles and nodes being the main sites of Zn localisation in the aerial parts. The highest Zn concentrations were observed in the epidermis and vascular system, of both leaves and stems, with distribution pattern differing in young and in mature leaves. This study highlights the Zn localisation patterns in S. plumbizincicola to improve our understanding of the mechanisms of hyperaccumulation. Growing in monoculture, S. plumbizincicola is an effective candidate for Zn phytoremediation in contaminated soils, with less promising results when co-cropped with the Ganges ecotype of N. caerulescens. Research Article Synchrotron elemental imaging reveals zinc distribution in the hyperaccumulator Sedum plumbizincicola (Crassulaceae) Julien Jacquet 1,2 , Ksenija Jakovljević 3 , Dennis Brueckner 4 , Catherine Sirguey 2 , Antony van der Ent 1,2,5 1 Econick SAS, F-54300, Lunéville, France. (https://orcid.org/0009-0009-1729-4053) 2 Université de Lorraine, INRAE, LSE, F-54000, Nancy, France. (https://orcid.org/0000-0002-3181-9014) 3 Department of Ecology, Institute for Biological Research Siniša Stanković - National Institute of the Republic of Serbia, University of Belgrade, Serbia. (https://orcid.org/0000-0002-1457-6807) 4 Deutsches Elektronen-Synchrotron DESY, 22607 Hamburg, Germany. (https://orcid.org/0000-0003-1714-5452) 5 Laboratory of Genetics, Wageningen University and Research, Wageningen, The Netherlands. (https://orcid.org/0000-0003-0922-5065) *Corresponding author: A. van der Ent ( [email protected] ) Abstract Sedum plumbizincicola is a zinc-cadmium (Zn-Cd) hyperaccumulator native to China with high potential for use in phytoremediation of contaminated soils in temperate climate. This study aimed to determine the Zn accumulation and distribution in S . plumbizincicola tissues grown on soils co-contaminated with Cd, Pb and Zn. The efficiency of Zn accumulation was assessed in a monoculture and in co-cropping systems with Noccaea caerulescens . The samples were analysed by inductively coupled plasma–atomic emission spectrometry and synchrotron micro-X-ray fluorescence elemental imaging. Sedum plumbizincicola grown in monoculture showed significantly higher foliar concentrations of Zn than the plants grown with N. caerulescens, with the leaf tips, petioles and nodes being the main sites of Zn localisation in the aerial parts. The highest Zn concentrations were observed in the epidermis and vascular system, of both leaves and stems, with distribution pattern differing in young and in mature leaves. This study highlights the Zn localisation patterns in S . plumbizincicola to improve our understanding of the mechanisms of hyperaccumulation. Growing in monoculture, S . plumbizincicola is an effective candidate for Zn phytoremediation in contaminated soils, with less promising results when co-cropped with the Ganges ecotype of N. caerulescens . KEYWORDS: Elemental localisation, phytoremediation, intercropping, Noccaea. INTRODUCTION Excessive elemental concentrations in the substrate induce various responses in plants, with exclusion being the most frequently observed defence mechanism (Bothe, 2011). However, in some plants, the concentrations of elements taken up by the roots and translocated to the aerial parts not only exceed those available in the soil, but are also 2–3 orders of magnitude higher than those typically found in non-accumulator species, classifying them as hyperaccumulators (Brooks et al., 1977; van der Ent et al., 2021). These concentrations were recognised as thresholds for hyperaccumulation and are elemental-specific (van der Ent et al., 2013). To date, hyperaccumulation of nickel (Ni) has been the most frequently observed (523 taxa; Reeves et al., 2018) and intensively studied in relation to Ni uptake, translocation and sequestration in the aerial parts of plants. In contrast, hyperaccumulation of zinc (Zn) was found in not much more than 20 taxa (Reeves et al., 2018), mainly in calamine soils characterised by excessive concentrations of cadmium (Cd), lead (Pb) and Zn (Wójcik et al., 2017). Interestingly, hyperaccumulation has also been reported in species growing in Zn-deficient soils where background Zn concentrations range between 10 and 100 μg g -1 . This suggests an affinity for Zn as a constitutive trait, as found in Noccaea caerulescens and Arabidopsis halleri , both model species for hyperaccumulation of multiple elements (Meyer & Verbruggen, 2012; Dinh et al., 2015; Stein et al., 2017; Merlot et al., 2021). Hyperaccumulation of Zn and Cd, as found in N . caerulescens and A . halleri , was also observed in two Sedum species, S . alfredii and S . plumbizincicola , a few examples outside the Brassicaceae family that hyperaccumulate these elements (Li et al., 2018). Sedum plumbizincicola is a relatively recently discovered hyperaccumulator, known only from the type locality in Zhejiang Province in China (Wu et al., 2013), with 14,600 μg g -1 Zn and 1470 μg g -1 Cd in shoots in its native habitat (Hu et al., 2015), and more than 18,000 μg g -1 Zn and 7000 μg g -1 Cd in shoots of hydroponically grown plants (Cao et al., 2014). The extremely high Cd and Zn concentrations in the leaves of S . plumbizincicola indicate a possible use for extraction within the principles of agromining, as at a biomass of 4–12 t ha -1 the removal efficiency in monoculture was estimated to be 215–515 g ha -1 Cd and 15–40 kg ha -1 for Zn (Deng et al., 2016). The multi-contamination tolerance of S . plumbizincicola , its perennial nature, rapid growth, high biomass production and its ability to be easily propagated vegetatively by cuttings, allowing 2‒3 potential harvests per year, add to the value of this process (Li et al., 2009; Hu et al., 2015; Wu et al., 2021; Song et al., 2022). In China, S . plumbizincicola has already been tested in monoculture under different climatic and edaphic conditions, however trials outside this area are extremely limited. According to the available literature, only one trial has been carried out in Europe under real conditions (Angelova, 2020). In this study, conducted in sub-alkaline soils (pH 7.7) in Bulgaria, a strong translocation potential for Cd, Pb and Zn (TF Cd and TF Pb > 2, and TF Zn > 4) was observed, exceeding the hyperaccumulation thresholds for Cd. The tolerance of S. plumbizincicola to water deficits and shade enables cultivation in co-planting systems with crops, such as wheat, maize, rice, sugarcane and cucumber (Zhao et al., 2011; Deng et al., 2016; Wu et al., 2021). Co-cropping is an agronomic strategy that is increasingly used together with crop density management practices, plant growth promoting bacteria and fungi, fertilisers and varietal selection to improve phytoremediation efficiency (Chaney et al., 2007; Kidd et al., 2015; Hossain et al., 2017; Bani et al., 2021; Benizri et al., 2021; Veerapagu et al., 2023; Wan et al., 2023). Phytoextraction efficiency in intercropping can be improved by mitigating the negative effects of the environment on hyperaccumulator species, by lowering pH and increasing elemental availability, or by overyielding when two hyperaccumulator species with different and complementary ecological niches are grown (Koelbener et al., 2008). For example, intercropping S. plumbizincicola with maize reduced the total concentration of Zn and Cd in the soil by 18.8% and 85.5%, respectively (Deng et al., 2016). In the remediation of multi- contaminated soils, the use of two (hyper)accumulating plant species can be particularly beneficial as the species enable complementary elemental accumulation when targeting different metals (Wang et al., 2022). However, the use of hyperaccumulators of the same elements may lead to accumulation with different efficiency, which is not only species-specific but may also strongly depend on the ecotype used (Jacquet et al., 2025). Considering the potential of S . plumbizincicola for agromining, this study aimed to investigate the accumulation and distribution patterns of Zn in shoots of plants grown under real field conditions (soils co-contaminated with metals) in temperate climates, i.e. outside the species native range, using synchrotron µXRF analysis, to better understand the mechanisms of hyperaccumulation. The accumulation capacity was additionally analysed in co-culture with N . caerulescens , another Zn hyperaccumulator species, to test different trends observed so far and to reveal patterns of Zn distribution. MATERIALS AND METHODS Plant culture conditions Seeds of N. caerulescens (Ganges ecotype, characterised in Gonneau et al., 2014) were collected in June 2019 in a former mining site in southern France and sown under greenhouse conditions in germination trays filled with horticultural compost. Cuttings of S. plumbizincicola were also grown under greenhouse conditions, from an individual collected at a contaminated site in China. The experiment was conducted under real field conditions in a contaminated urban garden in Forest-sur-Marque (near Lille, northern France). Analysis of the topsoil (0–20 cm), done prior to planting by ICP-OES (iCAP 6000 series, Thermo Scientific, Cambridge, UK), revealed moderate co-contamination with Zn (350 µg g -1 ), Pb (120 µg g -1 ) and Cd (1.5–2 µg g -1 ), with 13–40 µg g -1 of DTPA-extractable Zn (AFNOR, 1993), and a water pH of 6.0 to 6.5. Sedum plumbizincicola was grown in monoculture (Figure 1) at a density of 36 plants/m 2 for 10 months. To assess the effects of co-cropping on Zn accumulation and distribution, S. plumbizincicola was intercropped with N. caerulescens for the same period. In co-cultivation, S. plumbizincicola was planted at 8 plants/m 2 and N. caerulescens at 32 plants/m 2 . The experiment started in late June to early July 2022, with S. plumbizincicola directly planted as cuttings and N. caerulescens transplanted as seedlings of 12 weeks old. Harvesting and sampling took place twice, in September and in April of the following year. Sedum plumbizincicola was harvested only in the vegetative stage, while N. caerulescens was harvested either in the vegetative or flowering stage, depending on the plant. Bulk elemental analysis of plant samples After 3 and 10 months of cultivation, whole plants of N. caerulescens and S. plumbizincicola were collected from each treatment and bulked to make composite samples. Prior analysis, the aerial parts of the plants were carefully washed with tap water and rinsed with deionised water to remove soil dust and particles. The plant material was dried in an oven at 60°C for at least 48 h. Plant organs were ground to a fine powder (<200 µm) in an impact mill and weighed to 50 ± 5 mg in 15 mL polypropylene tubes. These samples were pre-digested with 1 mL HNO 3 (70%) and 2 mL H 2 O 2 (30%) for 16 h and then digested in a block heater (DigiPREP MS, SCP SCIENCE) for 3 h (ramping up and hold at 95°C). Samples were then diluted to 10 mL with ultrapure water (Millipore 18.2 MΩ cm -1 at 25°C) and filtered through 0.45 µm syringe filters before analysis by inductively coupled plasma–atomic emission spectroscopy (ICP-AES) using a Thermo Scientific iCAP PRO Duo X instrument. Synchrotron µXRF experiments The X-ray fluorescence microscopy experiments were performed at PETRA III (Deutsches Elektronen-Synchrotron DESY), a 6 GeV synchrotron radiation source, specifically at the hard X-ray microprobe undulator beamline P06 (Boesenberg et al., 2016). P06 is equipped with a cryogenically cooled double-crystal monochromator with Si (111) crystals. The X-ray beam can be focussed down to the sub-micrometre range using different focussing optics. An ion chamber upstream of the sample is used to monitor the incoming flux, while a 500 µm thick Si PIPS diode with an active area of 19 mm diameter (PD300-500CB, Mirion Technologies (Canberra) GmbH, Germany) downstream of the sample can be used to record the transmitted X-ray intensity in order to extract absorption data. Multiple XRF detectors enable the measurement of X-ray fluorescence data. The incident X-ray energy was 18 keV throughout the experiment and the beam was focused to 3.57 µm × 920 nm (h × v) using KB mirrors and prefocusing CRLs (compound refractive lenses), resulting in a flux of approximately 1.25 11 ph/s at the focus. For XRF detection, both a Vortex ME4 in 45° geometry and a prototype 16-element SDD Ardesia detector (800 µm thick chip with about 324 mm 2 combined active area for all 16 elements, Politecnico Milano, Italy (Utica et al., 2021)) in 315° geometry with Xspress 3 pulse processors were used. The fresh plant samples were brought from the experimental field in Forest-sur-Marque to DESY. The plant organs were sectioned by hand using a stainless steel razor blade (”dry knife method”), and mounted between two layers of Ultralene 4 μm thin foil to avoid evaporation, and immediately analysed. Data processing The XRF spectra were processed using non-linear least squares fitting as implemented in PyMCA (Solé et al ., 2007). After calibration using metal foils, this produced 32-bit .tiff files with pixel values corresponding to the µg cm -2 areal density of each element. The figures were prepared in ImageJ (Schneider et al., 2012) by changing the LUT to ’Fire’, adjusting the maximum values and adding concentration bars using the ‘calibration’ tool, and adding length scales. RESULTS Synchrotron µXRF analysis of the aerial part of S . plumbizincicola from monoculture revealed several important sites for Zn accumulation, with the highest concentrations observed in the leaf tips, petioles and nodes (Figure 2). Zinc accumulation in epidermal parts was observed in the leaf and stem cross-sections, with higher concentrations in the upper leaf epidermis, especially towards the tip. In the mesophyll, Zn concentrations were considerably lower and evenly distributed in the palisade and spongy tissues (Figure 3A). Besides the epidermis, most of the Zn was found in the vascular bundles of the stem, but only in alternating ones, while the others were depleted. Much lower concentrations of Zn were observed in the pith and cortex (Figure 3B). The large differences in foliar Zn accumulation in S . plumbizincicola were observed between plants from monoculture and those intercropped with N . caerulescens , but also between young and mature leaves (Figure 4). Mature leaves, especially those from the monoculture, showed the highest concentrations of Zn. The leaf tips, petioles (partially shown in Figure 4) and nodes were the main storage sites in all groups analysed, with preferential sequestration in vascular tissues in leaves with higher Zn concentrations. Unlike the vascular system, which showed an accumulation of Zn at high concentrations, localisation in the epidermis was found in all groups analysed, although with varying intensity. In co-culture with N. caerulescens , young leaves of S. plumbizincicola had significantly lower Zn concentrations in the vascular tissue than in the epidermis, whereas this effect was less pronounced in older leaves. The ICP analyses confirmed the differences observed by synchrotron µXRF between monoculture and co-cultivated S. plumbizincicola plants. In contrast to the 626 µg g -1 detected in plants grown in association with N . caerulescens for three months , foliar Zn concentrations in the S . plumbizincicola monoculture reached 1660 µg g -1 . In the N . caerulescens monoculture, Zn concentrations of 2980 µg g -1 were found in foliar tissues, whereas up to 4430 µg g -1 were detected in the leaves of the co-cultivated plants (Table 1). After 10 months, the monocultured plants of S . plumbizincicola accumulated more Zn than those in co-culture, with higher Zn concentrations observed – 3900 µg g -1 in the monocultured plants, and 1730 µg g -1 in the plants co-cultivated with N . caerulescens . However, in N . caerulescens , not only a decrease in foliar Zn concentration was observed at the end of the experiment, but also a reversal of the trend, as higher concentrations (2910 µg g -1 ) were observed in the monocultured plants compared to those grown in co-culture (1900 µg g -1 ). More subtle differences were observed in the concentrations of Cd and Pb in both species (Table 1), which are not discussed further, as Pb was below the detection limit for µXRF analysis and Cd could not be excited at the inference-free K-line at the incident energy used in this experiment. DISCUSSION The pattern of Zn localisation in S . plumbizincicola was found to be different in young and mature leaves and was mainly related to differences in elemental concentrations, which were much higher in the latter group. Whereas in plants grown with N . caerulescens , which had significantly lower foliar Zn concentrations, this element was accumulated in the epidermal tissues, in plants grown in monoculture where foliar Zn concentrations reached more than 5000 µg g -1 , Zn was evenly distributed both in the epidermis and in the vascular system (central and secondary veins). The predominant localisation of Zn in the epidermis is a common strategy in hyperaccumulator species (Scheckel et al., 2007; van der Ent et al., 2019, 2022), considering lower metabolic activity in these tissues, especially compared to the mesophyll. The epidermal tissues of the leaves and stems were found to be important for Zn localisation also in Sedum alfredii , another hyperaccumulator of Cd-Zn (Tian et al., 2009). In hydroponically grown S . plumbizincicola , however, the mesophyll was found to be almost equally important for Zn accumulation, especially in younger leaves with higher Zn concentrations, which is in contrast to the results of this study (Cao et al., 2014). The significant contribution of the mesophyll to the distribution of Zn was confirmed by the study of Hu et al. (2015), in which more than 50% of the Zn in mature leaves was found in this tissue layer. A similar finding was made for Zn in Arabidopsis hallerii (Küpper et al., 2000) and for Cd in Sedum alfredii (Tian et al., 2011), when the supply of these elements was high, but the epidermal cells and their vacuoles were not large enough to store the high metal concentrations (Küpper et al., 1999). In both hyperaccumulator Sedum species, the xylem also proved to be rich in Zn, indicating an efficient transport of Zn into the upper parts of the plant. In S . alfredii this enrichment was only observed in the hyperaccumulator ecotypes, but not in the non-accumulator ones (Tian et al., 2009), while the phenomenon of high Zn concentrations in alternating vascular bundles, which was also observed in S . plumbizincicola using the micro-PIXE technique (Hu et al., 2015) has not yet been clarified. High Zn concentrations in the nodes and petioles, preferentially in monocultured plants with higher Zn concentrations, additionally indicated efficient xylem loading and intensive translocation to the epidermal tissue, where it was predominantly sequestered. A similar pattern was also observed for Zn in Viola allchariensis (Jakovljević et al., 2023) and in Paulownia tomentosa (Azzarello et al., 2012), and for Cd in the Zn-Cd hyperaccumulator Potentilla griffithii (Qui et al., 2011). Intercropping, one of the most promising methods to improve metal extraction, showed contrasting results depending on the species or ecotype, even when two (hyper)accumulator species were used (Hu et al., 2019; Cao et al., 2021). When grown with N . caerulescens , a striking decrease in foliar Zn concentrations was observed in S . plumbizincicola compared to monocultured plants, with concentrations below the hyperaccumulation threshold (3000 µg g -1 ; van der Ent et al., 2013), although concentrations increased in with longer time of exposure. A similar finding, a strong decrease of Zn concentration in the shoot when co-cultivated with N . caerulescens , was observed in Salix dasyclados (Fuksová et al., 2009), indicating a higher efficiency of monoculture in phytoremediation practise. In N. caerulescens this trend largely depends on the exposure time, as after the initial predominant foliar accumulation of Zn, a reduction in concentration was observed in plants in co-culture, and after 10 months more Zn was accumulated in monoculture. Besides the stronger affinity of N . caerulescens for Zn uptake, which led to a depletion of available concentrations in the soil, this could also be due to its morphological characteristics, its well-developed root system and the better yield of aboveground tissue, making it significantly more competitive with S . plumbizincicola . Root exudates are another factor that strongly influences the efficiency of element uptake in co-cultures, mainly by altering pH and varying element availability, while a significant effect was found in the rhizospheric microbiome. Being species-specific and with abundance depending on soil elemental characteristics, bacterial community in soil are an important segment of phytoextraction strategy, that should be considered when planning co-cropping activities. Acknowledgements We acknowledge DESY (Hamburg, Germany), a member of the Helmholtz Association HGF, for providing experimental facilities. Parts of this research were carried out at PETRA III and we would like to thank Gerald Falkenberg and Jan Garrevoet for assistance in using beamline P06. The beamtime was allocated for proposal I-20220755 EC. This research was supported in part through the Maxwell computational resources operated at Deutsches Elektronen-Synchrotron DESY, Hamburg, Germany. The authors would like to thank the Pocheco (France) and Econick (France) companies for their collaboration in the Permagromine project. In particular, we acknowledge Guillaume Echevarria, Gabrielle Michaudel, Fanny Vautrin, Julien Verny, Pierre Vanden Berghe and their teams for growing the hyperaccumulators, maintaining the experiment and sampling. KJ acknowledges the support of Ministry of Science Technological Development and Innovation of the Republic of Serbia (Grant No 451-03-136/2025-03/ 200007). Author contributions JJ, CS and AvdE designed the study. JJ collected the samples, and JJ and CS collected and analysed the data. 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Agromining: Farming for Metals: Extracting Unconventional Resources Using Plants. Mineral Resource Reviews . (pp. 453–469). Cham: Springer doi:10.1007/978-3-030-58904-2_23 Wu, L. H., Liu, Y. J., Zhou, S. B., Guo, F. G., Bi, D., Guo, X. H., Baker, A. J. M., Smith, J. A., & Luo, Y. M. (2013). Sedum plumbizincicola XH Guo et SB Zhou ex LH Wu (Crassulaceae): a new species from Zhejiang Province, China. Plant Systematics and Evolution , 299(3), 487–498. doi:10.1007/s00606-012-0738-x Zhao, B., Shen, L. B., Cheng, M. M., Wang, S. F., Wu, L. H., Zhou, S. B., & Luo, Y. M. (2011). Effects of intercropping Sedum plumbizincicola in wheat growth season under wheat-rice rotation on the crops growth and their heavy metals uptake from different soil types. Chinese Journal of Applied Ecology , 22, 2725–2731. Table 1. Concentrations of Zn, Cd and Pb (in µg g -1 ) in foliar tissues of Sedum plumbizincicola and Noccaea caerulescens after three and ten months in monoculture and co-culture. Zn Cd Pb Zn Cd Pb Sedum plumbizincicola Monoculture 1660 6.4 0.5 3900 24.3 1.5 Sedum plumbizincicola Co-culture 626 9.6 0.3 1730 17.9 4.3 Noccaea caerulescens Monoculture 2980 102 0.2 2910 46.5 <DL Noccaea caerulescens Co-culture 4430 189 0.3 1900 47.3 0.4 Figure 1 . Sedum plumbizincicola after 10 months of cultivation on a moderately contaminated soil in the north of France. Figure 2 . Synchrotron µXRF elemental maps showing the distribution of Zn in shoots of Sedum plumbizincicola grown for 10 months in monoculture on a moderately contaminated soil . Figure 3 . Synchrotron µXRF elemental maps showing the distribution of Zn in cross-sections of [A] leaves and [B] stems of Sedum plumbizincicola grown for 10 months in monoculture on a moderately contaminated soil . Figure 4 . Synchrotron µXRF elemental maps showing the distribution of Zn in young leaves of Sedum plumbizincicola from monoculture [A] and co-cropping with Noccaea caerulescens [B], and mature leaves from monoculture [C] and co-cropping with Noccaea caerulescens [D] after 10 months of cultivation on a moderately contaminated soil. For easier comparison [A] and [B] have the same calibration bar, as do [C] and [D]. Four repetitions were performed for each treatment. Information & Authors Information Version history V1 Version 1 04 August 2025 Copyright This work is licensed under a Non Exclusive No Reuse License. Collection Ecological Research Keywords 39: chemical ecology 3: physiological ecology 51: material cycling biogeochemistry elemental localisation hyperaccumulator intercropping noccaea phytoremediation Authors Affiliations Julien Jacquet Botanickel View all articles by this author Ksenija Jakovljević 0000-0002-1457-6807 Institute for Biological Research "Siniša Stanković" National Institute of the Republic of Serbia, University of Belgrade View all articles by this author Dennis Brückner 0000-0003-1714-5452 Deutsches Elektronen-Synchrotron DESY, Germany View all articles by this author Catherine SIRGUEY 0000-0002-3181-9014 Université de Lorraine View all articles by this author Antony van der Ent 0000-0003-0922-5065 [email protected] The University of Queensland View all articles by this author Metrics & Citations Metrics Article Usage 259 views 170 downloads .FvxKWukQNSOunydq8rnd { width: 100px; } Citations Download citation Julien Jacquet, Ksenija Jakovljević, Dennis Brückner, et al. 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