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BTU
It is widely acknowledged that increasing complexity is a key attribute of ecosystem genesis. This is particularly true for primary succession on homogeneous substrates. However, a mechanistic understanding of spatial colonisation and pattern formation during primary succession has not been achieved yet. Thus, we studied this topic for 7 years within an experimental catchment (6 ha) established in the post-mining landscape of eastern Germany. Equidistant permanent plots (120 plots à 25m²) allowed for autocorrelation analyses, and thus tracing the spatial development of species cover performance. For each species tested, the “cover balance” first increased due to colonization, while decreasing in the course of succession. Drawing a benefit from these temporal trends, we suggested cover balance levels revealed by autocorrelation analyses to best indicate well-defined phases in primary succession at spatial entities. Hence, with the help of this spatial approach, terrestrial succession can be understood now much better at the most general level of the hierarchy provided by Pickett et al. (1987). In accordance with these authors, at subordinated levels of the causal hierarchy differential species performance might get collectively explained by contributing processes or conditions, such as ecophysiological traits, life history strategies, competition and allelopathy of the occurring plant species. Further research is needed to understand their relative contributions to the pattern formation.
Netzwerk Nagetier-übertragene Pathogene: Monitoring von Hantavirus-Infektionen in Deutschland
(2009)
In der Niederlausitz finden sich sehr unterschiedliche Beispiele für junge Bodenbildungen, die oftmals durch menschliche Eingriffe ausgelöst wurden. In dem Beitrag werden neben den jüngsten Bodenentwicklungen auf Bergbaufolgestandorten auch jüngere Böden auf seit dem Mittelalter mit Flugsanden überdeckten Standorten vorgestellt. Obwohl die Genese der auf diesen Flugsanddecken anzutreffenden Böden bereits seit bis zu 1 000 Jahre andauert, sind hier erst Böden der Klasse der Ah/C-Böden entwickelt. Auf den Bergbaufolgestandorten wird versucht, die Bodenbildung durch spezielle Meliorationsmaßnahmen zu fördern. Untersuchungen an diesen besonders jungen Böden lassen gut die Prozesse der initialen Bodenentwicklung erkennen.
Tree and shrub species grown on post-mining sites as source of biomass for bio-based industry
(2021)
Reclamation sites in East-Germany are characterized by very sandy substrates being almost free of organic matter, with extremely low nutrient contents and an annual precipitation of only 570 mm and lower. To enhance the value of such marginal lands for a future regionally based bioeconomy the EU funded BeonNAT project plans the plantation of regional tree and shrub species. The obtained forest biomass will be used for the production of biodegradable bio-based products and bioactive compounds that will play an important role to replace fossil-based competing substitute products. Field trials in different European countries will be established to test the benefit of intercropping/mixed-forest in marginal cultural land versus the natural growing/monoculture. In Germany, one test site is located in the post-mining sites of Lusatia. In this field trial the biomass development, the impact on soil fertility and biodiversity will be investigated after planting Robinia pseudoacacia and Rubus fructicosus.
In den 1980er Jahren wurde verstärkt die Frage diskutiert, ob archäologische Fundstücke aus Metall analog zu den damals beobachteten „neuartigen Waldschäden“ ebenfalls durch Immissionsbelastungen oder andere anthropogene Einflüsse gefährdet sind. Basis dieser Annahme waren Berichte von Archäologen und Restauratoren aus Museen und Denkmalpflege ganz Deutschlands, dass der Erhaltungszustand von neuen Eisen- oder Bronzefundstücken zunehmend schlechter werde. Um dem vermuteten Zusammenhang mit Umweltbelastungen nachzugehen, rief die Deutsche Bundesstiftung Umwelt Mitte der 1990er Jahre das interdisziplinäre Projekt „Schutz archäologischer Metallfunde vor immissionsbedingter Schädigung“ ins Leben. An verschiedenen archäologischen Fundstätten in der Bundesrepublik Deutschland wurden archäologische Metallfunde untersucht und parallel dazu Bodenuntersuchungen durchgeführt, um etwaige Umwelteinflüsse feststellen zu können. Die Ergebnisse
des Gesamtforschungsvorhabens sind in einem umfangreichen Band zusammengefasst1. Als ein mit Wald bedeckter Standort wurde der Döttenbichl, oder Döttenbühl, bei Oberammergau in die Untersuchungen mit einbezogen, auch weil vor allem die dort gefundenen Eisenobjekte unerwartet gut erhalten waren. Im Folgenden sollen die Ergebnisse der bodenkundlichen Untersuchungen an diesem Standort zusammengefasst werden, für die am 7. Mai 1996 Bodenproben genommen wurden.
Analyzing initial geomorphologic processes and structures: An alternative remote sensing approach
(2010)
Exploring hydrological and ecological processes plays a key role in understanding ecosystem development. In this respect, the constructed catchment, Chicken Creek (Hühnerwasser), has been established to promote fundamental and interdisciplinary scientific research. Since 2005, an on-going monitoring program has been established in this catchment to measure hydrological, biological, meteorological, and pedological parameters during the ecological development of the site. This comprehensive and multidisciplinary monitoring program has produced a large and diverse data set. Managing and exploring such a complex data set for research purposes can be a cumbersome task. As a consequence, we developed an online data portal https://www.b-tu.de/chicken-creek/apps/datenportal/ to efficiently handle the data from Chicken Creek catchment. The portal was constructed using R programming language, Shiny package, and the accompanying local Linux server. This platform allows for efficient data discovery, download, visualization, and analysis. Data visualizations are freely available, while data storage is limited to the authorized users. In comparison with the commonly used data base tools, R-Shiny offers several advantages. As an open source package, this platform allows for constructing a web data platform in an interactive way taking into account the complexity and diversity of the data. The flexibility of this routine enables to make an efficient user-demand data portal rather than relying on predetermined outputs. It also offers flexibility in data handling by using many different R packages, rendering versatile and extensive functionalities. R-Shiny is particularly powerful in terms of statistical analyses. Although the Chicken Creek online data portal is complete and available, new features, and extended capabilities are under active development. The Chicken Creek data portal provides a comprehensive and reliable database to give scientists a fast and easy access to all collected data.
Consequently, R-Shiny offers a great potential for future development of the web-based data portals to efficiently handle scientific data collected at different scales.
Marginal land (MagL) use for biomass production has received a lot of attention in the past decades. This alternative land use is free from conflicts with food crops, however it presents multiple challenges. The major issue to be resolved is the definition of marginal lands, which varies between disciplines and is often determined based on management goals. Moreover, concerns over the impacts of MagL use on environment, ecosystem services and sustainability have to be addressed.
In response to these issues, the SEEMLA approach proposes an algorithm and tools to support MagLs identification and mapping, as well as the assessment of their potential for biomass production. The SEEMLA algorithm assesses land marginality using the Muencheberg Soil Quality Rating (M-SQR) developed by Mueller et al. (2007). The algorithm also incorporates criteria to eliminate MagLs that may not be exploitable for bioenergy due to ecological and environmental issues or regulatory and legal restrictions and constraints posed by national or EU policies applied. Finally, MagL suitable for bioenergy production is categorized based on climatic zone and crop suitability. Alternative exploitation scenarios are developed for each class of MagL. Life Cycle Assessment (LCA) will determine the sustainability of each scenario, considering also the results of the pilot sites that have been established in Germany, Greece and Ukraine.
Experimental catchments with well-known boundaries and characteristics may contribute valuable data to hydrological, critical zone and landscape evolution research. One of the most well-established and largest constructed catchments is the Chicken Creek catchment (6 ha area including a 0.4 ha pond, Brandenburg, Germany) representing an initial ecosystem undergoing a highly dynamic ecological development starting from clearly defined starting conditions. The water balance dynamics of the catchment was calculated using a simple mass balance approach to reveal the impact of ecological development during 12 years. Water storage in the catchment was calculated from a 3D-model of groundwater volumes, soil moisture measurements and water level recordings of the pond. The catchment water balance equation was resolved for evapotranspiration, the only part that was not measured directly. Due to the known boundary conditions and the inner structure of the catchment, we were able to quantify the different storage compartments and their role in hydrologic response. Our results indicate that for small catchments with a highly dynamic ecological development like the Chicken Creek, the knowledge about saturated and unsaturated storage volumes enables a good estimate and closure of the water balance using a rather simple approach, at least in annual resolution. We found a significant relationship between vegetation cover in the catchment and calculated ET. Time series of meteorological, hydrological, soil and vegetation data over 12 years enabled us to characterize the transient development of the catchment and to evaluate the effect of different feedback mechanisms on catchment hydrology. The dataset from the Chicken Creek catchment indicate at least three phases in ecosystem development, where initial abiotic feedbacks (e.g. erosion) were followed by more and
more biotic controls (e.g. biological soil crusts, vegetation succession and growth). Data from Chicken Creek in high spatial and temporal resolution provide a valuable database underlining the high importance of abiotic/biotic feedback effects that change the hydrologic functioning and response of the catchment more than the water balance itself revealed and thus have to be included in catchment models.
Landscapes and ecosystems are complex systems with many feedback mechanisms acting between the various abiotic and biotic components. The knowledge about these interacting processes is mainly derived from mature ecosystems. The initial development of ecosystem complexity may involve state transitions following catastrophic shifts, disturbances, or transgression of thresholds. We propose a conceptual framework of feedback processes in early states of ecosystem development affected by spatiotemporal environmental drivers. To test this concept, we used 10-year time series of hydrological, biological, geomorphological, and soil data from the constructed catchment Chicken Creek.” The 6ha site was left to unrestricted development since 2005 and was intensively monitored. The data showed a very rapid development of the site with an increasing complexity and heterogeneity. In the first years, stochastic signals like the initial substrate conditions and external drivers like extreme weather events were the most important factors resulting in abiotic/abiotic feedback mechanisms shaping the morphology of the site and creating site diversity. Initial abiotic feedback mechanisms between water and substrate were soon followed by abiotic/biotic feedbacks between biological soil crusts, invading vegetation, geomorphology, and hydrology resulting in state transitions of catchment functioning.
Eine zunehmende Knappheit ackerbaulicher Standorte und der zugleich steigende Bedarf an nachwachsenden Rohstoffen für die energetische und stoffliche Nutzung lösen verstärkt Konflikte zwischen der Nahrungsmittelproduktion und der Biomasseproduktion aus. Marginale Standorte kommen entsprechend immer stärker in den Blickpunkt als mögliche Erweiterungsflächen für die Biomasseproduktion. In der wissenschaftlichen Auseinandersetzung fehlt allerdings eine eindeutige Definition für derartige marginale Standorte. Gerade im Hinblick auf eine mögliche Reduzierung von Nutzungskonflikten ist die Abgrenzung zu stillgelegten, aber agrarisch nutzbaren Standorten oder zu natürlichen Badlands, mit naturschutzfachlichen Potentialen oftmals schwierig. Das EU H2020-Projekt SEEMLA untersucht Potentiale und Verfügbarkeit von marginalen Standorten europaweit. In Deutschland, Griechenland und der Ukraine wurden Pilotstandorte eingerichtet, auf denen der Anbau von Bioenergiepflanzen exemplarisch durchgeführt wird. Für eine erste Bewertung der Standortspotentiale und auch der Ursachen der Marginalität der Standorte wurden Bodenuntersuchungen an allen Pilotstandorten nach den Vorgaben des Muencheberger Soil Quality Rating-Verfahrens (SQR) (1) durchgeführt und die Standorte klassifiziert. Das SQR-Verfahren bewertet ausgewählte Boden- und Standortseigenschaften und bezieht zusätzlich mögliche limitierende Faktoren (Hazard Indicators) wie Versauerung, Versalzung, Vernässung etc. mit ein, um eine Gesamtbewertungszahl zur Bodenfruchtbarkeit zu ermitteln. Dieser Beitrag stellt anhand der SEEMLA-Standorte exemplarisch Böden marginaler Standorte vor. Mit Hilfe der SQR Hazard Indicators werden charakteristische limitierende Faktoren vorgestellt. Es wird zudem versucht, die untersuchten Standorte in eine mögliche Klassifikation marginaler Standorte einzupassen und ihre Eignung für die Biomasseproduktion einzuschätzen.
The constructed catchment Chicken Creek was established in 2005 as an experimental landscape laboratory for ecosystem research. The 6 ha area with clearly defined horizontal as well as vertical boundary conditions was left for an unrestricted primary succession. The developing ecosystem offers the opportunity to study most processes occurring at the interface of bio-, pedo-, geo- and hydrosphere. Particularly interactions and feedbacks between different evolving compartments as well as the establishment of ecosystem functions and services can be investigated during the ongoing ecosystem development. The catchment is extensively instrumented since 2005 in order to detect transition stages of the ecosystem. Comprehensive data recorded with a high spatial and temporal resolution include hydrological, geomorphological, pedological, limnological as well as biological parameters.
This site offers the unique situation of an early stage ecosystem with highly dynamic system properties. The first years of development were characterized by a fast formation of geomorphological structures due to massive erosion processes at the initially non-vegetated surface. Hydrological processes led to the establishment of a local groundwater body within 5 years. In the following years the influence of biological structures like vegetation patterns gained an increasing importance. Feedbacks between developing vegetation and e.g. hydrological features became more and more dominant. As a result, different phases of ecosystem development could be distinguished until now.
This observatory offers manifold possibilities to identify and disentangle complex interactions between ecosystem compartments in situ under natural conditions. The originally low complexity of the system is growing with time facilitating the identification of influences of newly developing structures on ecosystem functions. Thus, it is possible to study effects of small-scale processes on the whole system at the landscape scale. In addition, the highly dynamic initial system properties allow the observation of multifaceted changes of ecosystem properties and functions within short periods of time.
We investigated the initial development of the landscape observatory ’Chicken Creek’, Germany, an artificial
catchment with well known boundary conditions and inner structures (Gerwin et al., 2011). Over a period of nine
years, we observed considerable changes within the site (Elmer et al., 2013). Both internal and external factors could be identified as driving forces for the formation of structures and patterns in the catchment. Over time,
secondary structures and patterns evolved and became more and more important. Invading biota and vegetation
succession initialized feedback mechanisms resulting in pattern and habitat formation as well as in increased differentiation, heterogeneity and complexity that are typical characteristics of ecosystems (Schaaf et al., 2013).
The processes and feedback mechanisms in the initial development of a new landscape may deviate in rates, intensity, and dominance from those known from mature ecosystems. It is therefore crucial to understand these early phases of ecosystem development and to disentangle the increasingly complex interactions between the evolving terrestrial and aquatic, biotic, and abiotic compartments of the system.
Within the context of severely disturbed landscapes with little or no ecological memory, such as post-mining
landscapes, we propose a simple framework that explains the catchment evolution as a result of competing feedbacks influenced by the initial conditions and the atmospheric drivers such as rainfall intermittency and intensity. The first stage of the evolution is dominated by abiotic feedbacks triggered by rainfall and subsequent fluid flow causing particle mobilisation on the surface and in the subsurface leading to flow concentration or in some instances to densification of surface and subsurface substrates. Subsequently, abiotic-biotic feedbacks start to compete in the sense that biological activity generally stabilizes substrate by preventing particle mobilisation
and hence contribute to converting the substrate to a habitat. We suggest that these competing feedbacks may
generate alternative stable states in particular under semi-arid and arid climatic conditions, while in temperate
often energy limited environments biological process “outcompete” abiotic processes leading to a stable state, in particular from the water balance point of view for comparable geomorphic situations.
The constructed catchment Chicken Creek was established in 2005 as an experimental landscape laboratory for ecosystem research. The 6 ha area with clearly defined horizontal as well as vertical boundary conditions was left for an unrestricted primary succession. All Critical Zone elements are represented at this site, which allows the
study of most processes occurring at the interface of bio-, pedo-, geo- and hydrosphere. It provides outstanding
opportunities for investigating interactions and feedbacks between different evolving compartments during ecosystem
development. The catchment is extensively instrumented since 2005 in order to detect transition stages of the
ecosystem. Data recorded with a high spatial and temporal resolution include ydrological, geomorphological,
pedological, limnological as well as biological parameters.
Conclusions
(2011)
Introduction
(2011)
Editorial for the Special Issue “Anthropogenic Geomorphology – Geomorphology in the Anthropocene”
(2023)
The cultivation of bioenergy plants at fertile, arable lands increasingly results in new land use conflicts with food production and cannot be considered as sustainable, therefore. Marginal lands have been frequently considered as potential alternative for producing bioenergy from biomass. However, clear definitions and assessment methods for selecting marginal lands and for calculating potentials are still widely missing. The project “SEEMLA” aims at triggering the exploitation of currently underused marginal lands for biomass production for energy purposes. Study sites have been selected in different European countries: Germany, Greece and Ukraine. The selected sites represent a wide variety of different types of marginal lands. Based on a soil assessment set given by the Muencheberg Soil Quality Rating (SQR) system potentially “marginal” sites have been investigated. The SQR system allows for clearly distinguishing between soils of higher and lower quality. Soils with SQR scores below 40 are regarded as “marginal”. They can be classified into different groups with regard to the importance of soil hazard indicators as valuated by the SQR approach. The calculated SQR scores correlate significantly with biomass yields of bioenergy plants. Further, the SQR method was adapted for use in a GIS study on marginal land potentials in Europe. 46 % of the investigated European area could be classified as “marginal” with SQR scores below 40. From that area 22.6 % can be considered as potentially suitable for producing renewable resources after eliminating protected sites or other places not suitable for any kind of land use. Taking the ecological demands of selected bioenergy plants into account it is possible to give first preliminary recommendations for regional crop cultivation. It can be concluded, that Europe offers a large potential for renewable resources from marginal sites. However, the implementation into practice is often impeded by missing or varying policies and regulation. A proper implementation needs clear regulations and also incentives for farmers at European level.
Severe disturbances of landscapes entail an ecosystem development with the formation of structures and functions which may reach either a new equilibrium state or a state similar to the original ecosystem. Natural disturbances can result from major events such as volcanoes, glaciers, or denudations from landslides. Major disturbance may also evolve from anthropogenic influences such as from mining operations. They all can be considered starting points for the development of ecosystems from ‘point zero’, which was one central research interest of Dieter Mueller-Dombois. In this paper results from research in the Lusatian post-mining landscapes (Eastern Germany) are presented. Different methodological approaches are discussed. The well-defined ‘point zero’ of the ecosystem development allows for research on chronosequence designs as well as real time series studies. Chronosequences have been investigated to gain insight into the medium to long-term direction of the development. Real time series are recorded to obtain a more detailed understanding. The paper is structured into three main parts: First, effects of ecosystem disturbances by mining in Lusatia and the initial conditions for restoration are presented. In the following part practical rehabilitation measures and land use options are discussed. Finally, the third part summarizes results of long-term monitoring in an artificial watershed. In conclusion, post-mining landscapes allow for relevant case studies of ecosystem development after severe disturbances. Particularly, the starting point of the initial phase is very well defined which distinguishes these anthropogenically disturbed landscapes from landscapes after natural disturbances.
After completion of the construction in 2005 (Gerwin et al. 2009), a major challenge was to develop and install a cross-disciplinary long-term monitoring program for the 6 ha area to record major environmental parameters adapted to the development of the site (Schaaf et al. 2013).
During its first 15 years, Chicken Creek showed a very dynamic development (Elmer et al 2013). Whereas the abiotic geosystem of the first 2-3 years was characterized by heavy erosion and sediment transport, primary succession by invading vegetation and the unexpected formation of soil crusts within few years resulted in more biotic-abiotic feedbacks that controlled catchment hydrology. Our observations over a period of 15 years indicate that even minor variations in initial substrate characteristics (e.g. texture) can have lasting impacts on geomorphical, hydrological and biological development like erosion intensity, groundwater levels or establishment of vegetation patterns. The time-series of monitoring data combined with a structure model of the catchment (Gerke et al. 2013) allowed the closure of the water balance by relatively simple calculations of water storage volumes and the estimation of evapotranspiration (Schaaf et al. 2017). From these data three stages of ecosystem development were derived which are characterized by increasing influence of biota and increasing system complexity.
Ecosystems are characterized by high complexity and are dynamically changing by abiotic and biotic drivers such as climate, vegetation and soil fauna. Due to feedback processes between compartments, ecosystems show an inherent degree of locally heterogeneous properties and structures at multiple spatial and temporal scales.
The constructed catchment “Chicken Creek” represents a unique and outstanding site to study an ecosystem at the initial stage of development. The catchment is located about 20 km south of the city of Cottbus (Germany). The 6-ha catchment area was built in 2005 of coarse-textured quaternary sediments from the adjacent lignite mine and forms a back- and foot-slope that flattens out to a pond. A clay liner of 2-3 m thickness seals the catchment at the bottom, allowing for the formation of a local water body fed by precipitation. After construction the site was left to an unrestricted and unmanaged succession to elucidate the role of structures und processes during the initial establishment of an ecosystem. To study processes of ecosystem development and newly emerging structures the Chicken Creek catchment is equipped with a comprehensive ecological monitoring network. A network of more than 40 wells and two weirs allow for recording groundwater levels, surface runoff and discharge. Meteorological parameters are recorded at three weather stations. Annual aerial photos and vegetation monitoring reveal the floral colonisation of the area. Here, we present the 12 years of diverse and extensive monitoring program which has been launched to measure hydrological, biological, meteorological, and pedological parameters during the ecological development of the catchment area.
The Huehnerwasser catchment was constructed 2004-2005 in a post-mining landscape in Eastern Germany and was left for an unmanaged primary ecosystem succession. At the same time a comprehensive monitoring program was launched for observing the expected dynamic development of this 6 ha site. During the following years this artificially created system was subject to a series of fast changes with regard to morphology, hydrology or vegetation cover. In summary, these processes resulted in significant alterations of the original, initial system within a short period of time. This development can be divided up into single phases with dominating processes or groups of processes. A first phase was mainly characterized by interactions between abiotic system components such as water and substrate. However, already in this very early developmental phase biotic components in the form of microorganisms were a crucial system forming factor by means of biological soil crusts. This period was quickly replaced by a second phase with dominating hydrological processes. During this phase the local groundwater body of the system developed to its full extent. With the beginning of the present phase biota gained control of the system behavior. Particularly plants and related processes such as water uptake and evapotranspiration are now responsible for new modifications of the system. The Huehnerwasser site offers an example of young and highly dynamic ecological systems. This transitional development of a very young ecosystem contrasts with the behavior of mature ecosystems with slow and less pronounced changes unless the system is disturbed. In this paper we present insights into a unique landscape observatory which illustrates the dynamic nature of natural systems in fast motion.
Der Lehrstuhl Bodenschutz und Rekultivierung der Brandenburgischen Technischen Universität Cottbus-Senftenberg (BTU CS) ist Projektpartner in dem von der Europäischen Union im Rahmen des Programms Horizon 2020 geförderten Forschungsprojekts SEEMLA → „Sustainable exploitation of biomass for bioenergy from marginal lands in Europe“ → „Nachhaltige Nutzung von Biomasse für Bioenergie von marginalen Standorten in Europa“. Übergeordnetes Ziel des Projektes ist die Ermittlung des Biomassepotenzials, das auf marginalen Standorten innerhalb von Europa nachhaltig produziert und energetisch verwertet werden kann. Die BTU CS hat in diesem Kontext Untersuchungen zur Bewertung der Marginalität von potenziellen Biomasse-Produktionsstandorten in Deutschland, Griechenland und der Ukraine durchgeführt. Zu diesem Zweck wurde das Müncheberger Soil-Quality-Rating System (SQR) (Müller et al. 2007) eingesetzt.
Our investigations at the artificial catchment ´Chicken Creek´ in Lusatia/Germany aim to disentangle and understand the feedback mechanisms and interrelationships of processes and their co-development with spatial and temporal structures and patterns by studying this initial, probably less complex ecosystem. Intensive measurements were carried out in the catchment with regard to the development of surface structures, hydrological patterns, and vegetation dynamics.
During the first seven years, considerable changes within the catchment were observed. Both internal and external factors could be identified as driving forces for the formation of structures and patterns in the artificial catchment. Initial structures formed by the construction process and initial substrate characteristics were decisive for the distribution and flow of water. External factors like episodic events triggered erosion and dissection during this initial phase, promoted by the low vegetation cover and the unconsolidated sandy substrate.
The transformation of the initial geo-system into areas with evolving terrestrial or aquatic characteristics and from a very episodic to a more permanent stream network and discharge, together with the observed vegetation dynamics increased site diversity and heterogeneity with respect to water and nutrient availability and transformation processes compared to the more homogenous conditions at point zero.
The processes and feedback mechanisms in the initial development of a new landscape may deviate in rates, intensity and dominance from those known from mature ecosystems. It is therefore crucial to understand these early phases of ecosystem development and to disentangle the increasingly complex interactions between the evolving terrestrial and aquatic, biotic and abiotic compartments of the system. Artificially created catchments could be a suitable tool to study these initial developments at the landscape scale under known, designed and defined boundary conditions.