Frontiers | Lacto-fermented vegetables as dynamic food ecosystems: emerging mechanisms along the microbiome–gut

Lacto-fermented vegetables as dynamic food ecosystems: emerging mechanisms along the microbiome–gut–brain axis

• 1. Charité Competence Center for Traditional and Integrative Medicine (CCCTIM), Charite – Universitatsmedizin Berlin, Berlin, Germany

• 2. Department of Biotechnology, Technische Universität Berlin, Berlin, Germany

• 3. Institute of Environmental Biotechnology, Graz University of Technology, Graz, Austria

• 4. Department of Microbiome Biotechnology, Leibniz Institute for Agricultural Engineering and Bioeconomy, Potsdam, Germany

• 5. Department of Pediatrics, University of São Paulo, São Paulo, Brazil

Abstract

Lacto-fermented vegetables (LFVs) are traditional foods produced through the microbial transformation of plant materials by lactic acid bacteria, with additional contributions from yeasts and other microorganisms. Fermented foods have been increasingly investigated for their potential modulation of the microbiome-gut-brain axis. This research has mainly focused on dairy fermentations, leaving LFVs largely underexplored. This mini review examines LFVs and their emerging mechanisms within the microbiome-gut-brain axis (MGBA). It emphasizes their character as evolving microbial ecosystems and as sources of bioactive metabolites. During the fermentation process, ecological succession drives the production of a variety of compounds, including organic acids, amino acids and lipid derivatives, transformed phytochemicals, structural microbial components, and micronutrients. Many of these compounds have plausible connections to MGBA routes. These links include modulation of the intestinal barrier, local and systemic immune responses, enteroendocrine signaling, and autonomic/vagal pathways. Particular focus is given to the small intestine as an early signaling interface between metabolites and host physiology. Scientific evidence of the beneficial effects of LFVs on human health is limited. Existing studies show improvement in gastrointestinal symptoms, cardiometabolic risk factors and gut microbial function. Studies connecting LFVs to neurological and psychological health are lacking. We propose understanding LFVs as complex food ecosystems whose effects on host physiology arise from interactions among microorganisms, metabolites, environmental conditions, and host responses. Future research integrating food multi-omics, microbiome analyses, immune profiling, neuroendocrine measurements, and clinical outcomes may help clarify the role of LFVs in MGBA and support the development of targeted fermentation-based nutritional strategies for intestinal, immune, and neuropsychological health.

1 Introduction

Fermented foods are characterized by the desired growth of microorganisms transforming the food matrix over time (). Food fermentation has been used by humans for millennia (, ). Campbell-Platt et al. estimated the proportion of fermented foods worldwide in 1994 at around one-third of overall food consumption (). The benefits of preservation, increased food safety and digestive efficiency, and, in some cases, alcohol content likely contributed to the early use of fermentation technologies (, ). A large variety of foods are fermented all around the world, including dairy, fruits, vegetables, leaves, cereals, tubers, meat, fish and mushrooms, involving an even larger variety of specialized microorganisms ().

Despite this diversity, common principles of community assembly can be identified across fermented foods (, ). Community composition is determined by dispersal (how microorganisms reach an environment), selection (environmental pressures), diversification and ecological drift, with dispersal and selection being especially useful in the context of food fermentation (, ). In fermented foods, dispersal includes the resident microorganisms present in the raw food, the surrounding environment, as well as starter cultures. Previous work has, for instance, shown that different crop management practices can shape the plant microbiome (). Temperature, salinity, acidity, oxygen availability, nutrient composition, and the activity of bacteriophages, yeasts, and fungi are important environmental selection factors in fermented foods (, , ). Interestingly, selective pressures and substrate seem to outweigh geographic influences, as similar microbial communities are regularly observed in comparable fermented products across several continents (, ).

Existing reviews have detailed how fermented foods serve as carriers for beneficial microbes and metabolites and can modulate the central nervous system (–). Several reviews also focused on the potential of fermented foods to moderate neurologic and psychiatric health conditions such as depression and anxiety or neurodegenerative diseases (, , )

Until now, most research linking fermented foods to the MGBA has focused on dairy ferments such as yogurt and kefir. Lacto-fermented vegetables (LFVs) offer a distinct profile characterized by plant- and soil-derived starter cultures and the development of plant-derived bioactive metabolites, including transformed phytochemicals and dietary fiber (, ). Throughout diverse vegetable fermentations, the microbial communities are typically dominated by lactic acid bacteria (LAB), particularly species of the genera Lactobacillus, Leuconostoc, Weissella, Pediococcus, and Lactococcus (, , ).

This mini review will focus specifically on LFVs and their possible mechanisms of action on the central nervous system (CNS). The particular perspective centers around an understanding of LFVs as complex food matrices that may act dynamically over time through combined direct and indirect pathways (Figure 1).

2 Biochemical evolution in LFVs

2.1 Microbial succession

The transformation of a raw vegetable into bioactive fermented foods is controlled by a time-dependent microbial and biochemical succession (, ). Lacto-fermentation occurs in a low-oxygen environment where salt, acidification, substrate composition and microbial interactions select for specialized LAB (). Initial communities derive largely from plant- and soil-associated microorganisms. Although LAB may occur at low abundance before fermentation, their presence and succession can be influenced by geographical region, soil properties, farming practices, plant genotype and post-harvest handling (). Spontaneous LFV fermentation may therefore retain microbial signatures from the agricultural ecosystem, including non-LAB members, before conditions select for acid-tolerant LAB (). This links LFVs to a broader soil-plant-food-microbial continuum (, , ).

Early fermentation is frequently dominated by heterofermentative species such as Leuconostoc mesenteroides and Weissella spp., producing lactate, acetate, CO2 and ethanol. As pH decreases, communities typically move toward more acid-tolerant homofermentative LAB, including Lactiplantibacillus plantarum and related Lactobacilli (, ). This ecological succession shapes the developing metabolic landscape of LFVs (–).

2.2 Organic acids

Lactate is one of the most abundant metabolites in many LFVs (, , ). It drives acidification, supporting pathogen suppression (, ). Beyond preservation, orally administered lactate has been linked to enteroendocrine effects in humans, including increased GLP-1, insulin, and glucagon, reduced ghrelin, and earlier satiety (, ). Lactate can also activate the human HCA1 receptor, which is involved in intestinal barrier integrity and inflammatory signaling (). In preclinical and in vitro models, lactate exhibited barrier-protective and anti-inflammatory effects and supported the expansion of beneficial taxa such as Akkermansia muciniphila, with downstream effects on regulatory T cells (, , ).

Acetate is produced by LAB and may also arise through cross-feeding (, ). It is rapidly bioavailable after ingestion and has been associated with improved fasting glucose in humans (, ). Acetate activates the short-chain-fatty acid (SCFA) receptors GPR43 and GPR41, involved in immune regulation, gut hormone synthesis, barrier integrity, and neuronal function (). Other SCFA, including propionate and butyrate, may also be present in some LFVs or arise after fermentation-derived compounds are further metabolized by resident microbes (31–34).

2.3 Amino-acid and lipid-derivatives

LAB can convert amino acids into active metabolites (35). Phenylalanine can be transformed into D-Phenyllactate (D-PLA), which increases in plasma and urine after sauerkraut consumption and acts as a potent agonist of the human HCA3 receptor (36–38). Tryptophan-derived indole-3-lactic acid (ILA), detected in different LFVs, can activate HCA3 and the Aryl hydrocarbon receptor (AhR), a transcription factor involved in barrier regulation and immune signaling (, 39–41). Interestingly, the human-specific HCA3 receptor may represent an evolutionary adaptation to the habitual consumption of fermented foods, converting microbial signals into immune responses via monocytes and the intestinal epithelium (, 38). LAB-driven decarboxylation of glutamate can generate gamma-aminobutyric acid (GABA), an inhibitory neurotransmitter (42). Although orally ingested GABA likely has limited direct access to the brain, peripheral GABAergic signaling may influence gut motility, secretion, inflammation and immune tolerance (, 43, 44). LFVs can also contain other amino acid derivatives, such as 2-hydroxyisocaproic acid and 12,13-DiHOME, with potential relevance to microbial signaling, barrier and immune interactions (45–47).

2.4 Transformed phytochemicals

LFVs contain plant secondary metabolites, typically derived from the raw plant (, 48). LAB enzymes, including ß-glucosidases, can convert glycosides into smaller aglycones with potentially higher bioavailability (49–51). In cruciferous vegetables, fermentation promotes glucosinolate degradation and the formation of indole-derivatives such as indole-3-carbinol and ascorbigen, which may contribute to antioxidant and anti-inflammatory properties (52, 53). Fermentation may also increase total phenolic content by releasing bound phenolic derivative compounds (, 54). Some phenolic compounds and their metabolites have been associated with metabolic, cardiovascular and neurologic benefits, although LFV-specific human data remain limited (, 55–58).

2.5 Bacteriocins, exopolysaccharides and microbial structural components

LAB produce bacteriocins, antimicrobial peptides that can shape microbial ecology by suppressing competing bacteria (, 59–61). Exopolysaccharides are another class of fermentation-associated bioactives and may support barrier integrity, community structure and mucosal immune modulation (, , 62, 63).

LFVs also contain non-l fragments, lipoteichoic acids, CpG DNA and surface proteins (64–66). These microbe-associated molecular patterns (MAMPs) interact with host pattern recognition receptors (PRRs). Those interactions are context-dependent: in balanced settings, microbial sensing can support immune training, whereas in dysregulated states it may drive inflammatory responses (, 67)

2.6 Micronutrients and reduction of anti-nutrient

Fermentation can modify the micronutrient profile of vegetables and fruits. LFV may show increased concentration of B vitamins, including folate, riboflavin, vitamin B12 and vitamin K (, 68, 69). Effects on vitamin A and vitamin C vary by substrate and fermentation (, 68, 70). Although some mineral concentrations may decline (e.g., iron, magnesium, calcium), mineral bioavailability can improve through microbial phytase activity, which reduces phytates that otherwise bind cations (, 71).

2.7 Bacteriophages and yeast in LFVs

LFVs contain bacteriophages that co-evolve with bacterial communities and shape succession by infecting LAB and other fermentation bacteria (, 72, 73). Phage-mediated lysis limits overgrowth and releases microbial material that supports cross-feeding (74, 75). Yeast, including Saccharomyces spp. and Candida spp., also participate in carbohydrate metabolism, flavor formation, oxygen availability, nutrient synthesis and microbial succession, thus affecting the functional properties of LFVs (, 76, 77).

3 Direct and indirect modulation of the MGBA: overlapping routes

The microbiome communicates with the brain through bidirectional routes collectively termed the microbiota-gut-brain axis (MGBA) (78, 79). Known mechanisms include changes in gut microbiome ecology, immune and inflammatory signals, intestinal and blood-brain barrier integrity, the hypothalamus-pituitary-adrenal (HPA) axis, neurotransmitters, gut hormones and vagal signaling (78–80).

In the context of LFVs, direct and indirect MGBA effects should be viewed as overlapping rather than separate pathways (Table 1). LFVs deliver microbial components and fermentation-derived metabolites that interact with intestinal barrier, immune, endocrine and neuronal interfaces. Additionally, they provide live-plant-associated microorganisms that can transiently engage with resident communities via cross-feeding and competition (, , , 79, 81). Plant-associated microorganisms specific to LFVs may therefore contribute indirectly to MGBA-relevant pathways by modifying the microbial and metabolic starting conditions of fermentation (, ).

LFV-derived bioactive components and plausible MGBA-relevant routes.

At the CNS level, the most plausible LFV-related mechanisms are indirect neuroimmune and neurovascular pathways. Microbial metabolites, such as SCFAs, regulate microglial maturation, homeostasis and function, linking intestinal microbial metabolism and CNS innate immune tone (81, 82). Gut microbiota and SCFA have also been implicated in blood-brain-barrier integrity (81, 83). Although these mechanisms are not LFV-specific, they offer a framework through which LFV-derived organic acids, cross-feed-derived SCFAs, indole derivatives and microbial structural components may influence brain-relevant physiology (84).

LFVs and other fermented foods deliver live LAB and associated microbial communities, usually resulting in transient colonization (, 85). These transient communities interact with resident microorganisms through cross-feeding or competition (, 86). Studies have reported variable effects of fermented-food-rich diets on gut microbial diversity (87–89). Additionally, viable and non-viable microbial components can act as MAMPs recognized by the immune system, possibly supporting immune tolerance or inflammation, depending on context (66, 90).

Several studies suggest that metabolites produced during lactic acid fermentation may be sufficient to drive gut immune and barrier effects, even without live microbes (). In Caco-2 cell barrier models, the complete fermented cabbage matrix protected against cytokine-induced barrier damage better than single metabolites such as lactate, D-PLA and ILA, and these effects were not strictly dependent on microbial survival (47). Similarly, fermented vegetable brine and microbe-free supernatants showed comparable microbiota restructuring and barrier-tolerogenic activity (). In patients with irritable bowel syndrome, unpasteurised and pasteurized sauerkraut led to similar symptom improvement (91). LFV consumption may also alter fecal metabolites more strongly than microbial composition, supporting functional gut microbiome modulation without durable colonization (92).

A common immune-mediated MGBA pathway is the modulation of mucosal barrier function and systemic inflammatory tone. Gut microbiota and microbial metabolites can influence mucosal immunity and systemic cytokine milieu, thereby affecting brain function and behavior (84). In humans, a high-fermented-food diet reduced multiple inflammatory markers while increasing microbiota diversity, although the intervention was not strictly LFV-specific (88). In preclinical work, fermented vegetable brine promoted tolerogenic barrier immunity, implying reduced pro-inflammatory signaling capacity from the gut ().

Barrier modulation represents a closely related pathway (71). Strengthening epithelial tight junctions and the mucus layer can reduce the translocation of inflammatory luminal products, thereby altering systemic immune signaling (). Fermented foods can strengthen intestinal barrier through metabolites, including SCFAs and organic acids (, , , 93). LFVs provide lactate and fermentable plant substrates that resident microbes convert into SCFAs, including butyrate and propionate (). SCFAs regulate barrier function and mucosal immune tone (94, 95). Propionate can increase satiety via gut hormones such as PYY and GLP-1, while butyrate supports colonocyte energy metabolism and epithelial integrity (94, 96).

LFVs are also discussed as food sources of neuroactive microbial metabolites (, , ). One example is gamma-aminobutyric acid (GABA), produced by LAB via glutamate decarboxylation (42). Orally ingested GABA has limited direct access to the brain (97). However, LFV-derived GABA can participate in peripheral enteric, immune and inflammatory signaling, including effects on gut motility and secretion (43). Thus, LFV-derived GABA should be interpreted as a peripheral neuroimmune and enteric signaling molecule, not as a direct CNS neurotransmitter (97, 98). Visceral signaling then propagates to the CNS indirectly via immune, endocrine and neural routes, including vagal afferents ().

Changes in bowel habits and visceral sensitivity are MGBA-relevant because disorders of gut-brain interaction, such as irritable bowel syndrome (IBS), involve motility changes, visceral signaling, enteric nervous system activity, microbiota-immune interactions, and CNS processing (99). Sauerkraut consumption improved symptom severity in patients with IBS (91). While this was not a direct brain-outcome study, improvement in IBS-related symptoms supports the relevance to gut–brain sensory and immune signaling, including spinal and vagal afferent pathways (80, 99).

Vegetable fermentation also transforms phytochemicals and plant matrices, generating polyphenols and other small bioactive metabolites (, 100). Some polyphenols and microbial metabolites can enter systemic circulation and may interact with the blood-brain barrier (101, 102). Evidence from broader polyphenol research suggests that selected polyphenols and their metabolites can cross the blood-brain barrier, modulate microglia and influence neurotrophic pathways, such as BDNF-related signaling involved in synaptic plasticity (, 103–106). Experimental MGBA studies further show that individual microbial metabolites can have brain-region- and cell-type-specific effects in preclinical models (107). These data support metabolite-specific gut-brain mechanisms, although they are not yet LFV-specific.

Overall, direct evidence for LFV effects on the MGBA remains limited. While fermented foods, the gut microbiome, and psychobiological outcomes are increasingly investigated, LFV-specific mechanistic and clinical data remain scarce. Links between LFVs and MGBA-related outcomes should therefore be interpreted as mechanistically plausible routes deduced from broader fermented foods and MGBA research. Targeted mechanistic and human interventional studies are needed to clarify these associations.

4 The importance of the small intestine and enteroendocrine cells

Most human microbiome studies use fecal samples, which mainly reflect distal colonic conditions (108). However, food-derived microbes and metabolites travel through the mouth, stomach, duodenum, jejunum and ileum before reaching the colon. The small intestine is likely an important signaling site that differs markedly from the colon in nutrient availability, pH, oxygen exposure, transit time, immune architecture and enteroendocrine activity (109, 110). The ileum holds highly active immune monitoring structures such as Peyer’s patches (111). It is likewise enriched in enteroendocrine signaling related to motility, appetite control and gut-brain communication (110, 111).

Enteroendocrine cells sense luminal nutrients, microbial metabolites and bile acids through G-protein-coupled receptors, including GPR41, GPR43, HCA-receptors and bile acid receptors. (112–114). They translate these signals into endocrine, paracrine, and neural pathways, releasing mediators such as GLP-1, PYY, cholecystokinin, serotonin, and ghrelin, thereby affecting appetite regulation, motility, secretion, glucose metabolism, autonomic activity, visceral signaling, and stress-related pathways (113–116). Specialized enteroendocrine subtypes, termed neuropod cells, form synapse-like connections with enteric neurons and vagal afferents, allowing fast transmission of luminal signals to neural pathways (112, 115–118).

These mechanisms may be relevant for LFVs because fermentation-derived metabolites may be sensed before extensive colonic transformation occurs (86). Spencer et al showed that sauerkraut brine altered fecal and caecal microbial composition, with the strongest changes observed in the ileum, where Akkermansia muciniphila and regulatory T cells were enriched (). These data support the plausibility that LFV-derived compounds may influence enteroendocrine, neural, immune and barrier-related pathways before reaching the colon. However, direct human evidence for LFV-signaling in the small intestine is still limited.

5 Health outcomes of LFVs in humans

Existing reviews and intervention studies indicate possible benefits for gastrointestinal health, cardiometabolic risk factors and gut microbial function (, ). However, no human study has directly connected LFV-consumption to MGBA-measures. In patients with irritable bowel syndrome, daily sauerkraut consumption for six weeks improved symptom severity and altered gut microbial composition, with no significant difference between pasteurized and unpasteurised products (91). In healthy adults, LFV effects on gut microbial composition appear product- and person-dependent. A sauerkraut cross-over trial detected changes in individual bacterial species and heightened serum SCFA after pasteurized sauerkraut consumption (119).

Evidence from Kimchi studies is more extensive. A scoping review including 11 RCTs found ameliorations in serum lipid profiles and body-fat-related parameters (120). A systematic review and meta-analysis of intervention and prospective cohort studies found improvements in fasting glucose, triglycerides and blood pressure, with observational evidence linking higher kimchi intake to normal weight and lower risk of metabolic syndrome and cancer (121).

These preliminary findings support the health-improving effects of LFVs. Nevertheless, human studies linking LFVs to neurologic and psychological health are still scarce. Ongoing clinical research is beginning to address this gap by integrating microbiome, immune and patient-reported outcome measures (PROMs). A recently published RCT protocol investigates fermented vs. unfermented red cabbage effects on allergic rhinitis symptoms, immune markers, the gut microbiome, and PROMs for depression and anxiety in young adults (122).

On a final note, we would like to emphasize that LFVs can show considerable microbial and compositional variability, which may sometimes raise safety concerns for human consumption. Surveys of homemade and commercial LFVs generally found mature products free of major pathogens. However, considerable heterogeneity in acidity, LAB and yeast abundance, and salt content was observed, with occasional detection of potentially undesirable microorganisms (123, 124). In a challenge study in which pathogens where inoculated into sauerkraut, they remained viable during early fermentation, particularly when acidification was delayed (125). LFVs may also contain biogenic amines and high salt concentrations, which may pose further challenges for human consumption (126). Good hygiene and well-controlled fermentation conditions are therefore essential.

6 Future directions

Future human studies should connect LFV-mediated microbial and metabolic changes with immune, neuroendocrine and clinical outcomes. Relevant MGBA-endpoints may include intestinal barrier markers, salivary cortisol as HPA-axis marker, heart-rate variability as an autonomic proxy, neuroimmune cytokine panels, and neurocognitive and affective outcomes, including neuroimaging techniques. Additionally integrating food multi-omics could help identify LFV preparations with specific mechanistic properties. This direction is consistent with the concept of engineered LAB-fermentates as functional fermented foods with defined health-supporting properties (79, 81, 127).

Research on specific bacteria, bacteriocins, or bacteriophages in LFVs may yield important insights into pressing current problems such as antimicrobial resistance.

Another research avenue is to explore how agricultural methods influence LFV composition and possible health effects. Different cultivation systems shape plant-associated microbiomes. Raw plant material entering fermentation carries microbial signatures influenced by farming practices. This links soil, plants, plant fermentation, and the human gut microbiome ().

LFVs may also serve as controllable complex model ecosystems for studying food-gut microbiome-host interactions. Fermented foods are dynamic ecological systems in which bacteria, fungi, bacteriophages, metabolites and environmental conditions interact, giving rise to emergent functions. Upcoming research could leverage this property to explore how external pressures, such as cultivation practices, fermentation conditions, or antibiotic exposure, influence ecosystem evolution, resilience, invasion resistance, recovery after perturbation, and transition toward dysfunctional states (, 128).

Overall, LFVs remain an intriguing but underexplored area within fermented-food research. Future advances in our understanding of LFVs may help clarify their potential as targeted components of nutritional strategies for intestinal, immune and neuropsychological health.

Statements

GN: Conceptualization, Data curation, Investigation, Methodology, Project administration, Validation, Visualization, Writing – original draft, Writing – review & editing. SN: Writing – review & editing. FN: Writing – review & editing. AD: Writing – review & editing. NC: Writing – review & editing. CB: Writing – review & editing. WW: Writing – review & editing. GB: Supervision, Writing – review & editing. GS: Re

Funding

The author(s) declared that financial support was received for the publication of this article. The article processing charge was covered in part by the Open Access Publication Fund of Charité – Universitätsmedizin Berlin and in part by Ekhagastiftelsen, grant number 2024-155.

Conflict of interest

The author(s) declared that this work was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

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Abbre

5-HT, 5-hydroxytryptamine (serotonin); AHR, Aryl hydrocarbon receptor; BBB, Blood–brain barrier; BDNF, Brain-derived neurotrophic factor; CCK, Cholecystokinin; CNS, Central nervous system; CpG DNA, Cytosine-phosphate-guanine; DNAD-PLA, D-phenyllactate; EEG, Electroencephalogram; EEC, Enteroendocrine cell; ENS, Enteric nervous system; FF, Fermented foods; fMRI, Functional magnetic resonance imaging; GABA, Gamma-aminobutyric acid; GLP-1, Glucagon-like peptide-1; GPCR, G protein-coupled receptor; HICA, 2-hydroxyisocaproic acid; HPA, Hypothalamic–pituitary–adrenal (axis); IBS, Irritable bowel syndrome; ILA, Indole-3-lactic acid; LAB, Lactic acid bacteria; LFV, Lacto-fermented vegetables; MAMP, Microbe-associated molecular pattern; MGBA, Microbiome–gut–brain axis; PRR, Pattern recognition receptor; PROM, Patient-reported outcome measure; PYY, Peptide YY; RCT, Randomized controlled trial; SCFA, Short-chain fatty acids; Treg, Regulatory T cells.

References

Summary

bioactive metabolites, fermentation, fermented foods, gut microbiome, lactic acid bacteria, lacto-fermented vegetables, microbiome-gut-brain axis

Ngoumou GB, Ngandeu Schepanski S, Nejati F, Diedering A, Cassagnau N, Beyrow C, Wicaksono WA, Berg G and Seifert G (2026) Lacto-fermented vegetables as dynamic food ecosystems: emerging mechanisms along the microbiome–gut–brain axis. Front. Nutr. 13:1903488. doi: 10.3389/fnut.2026.1903488

Marta Maria Nowacka-Chmielewska, Jerzy Kukuczka Academy of Physical Education in Katowice, Poland

Rasha A. F. Jasim, University of Babylon, Iraq

Jelang Jelku D. Sangma, Central Agricultural University (Imphal), Meghalaya, India

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ORCID: Gonza B. Ngoumou orcid.org/0000-0001-6501-732X; Steven Ngandeu Schepanski orcid.org/0000-0002-7695-1289; Fatemeh Nejati orcid.org/0000-0002-8991-7442; Alina Diedering orcid.org/0009-0001-2595-7090; Wisnu Adi Wicaksono orcid.org/0000-0002-1556-1981; Gabriele Berg orcid.org/0000-0001-9423-3101; Georg Seifert orcid.org/0000-0002-7109-9277