Cited by 374Bacterial biofilms and quorum sensingWastewater Treatment and Nitrogen RemovalPolysaccharides Composition and Applications

The DOI matches its Crossref record; 96 of the 96 references it cites were found in a registry or an index.

EPS—Then and Now

Open access
Flemming, Hans-Curt·2016·Microorganisms

Abstract

“Slime” played a brief and spectacular role in the 19th century founded by the theory of primordial slime by Ernst Haeckel. However, that substance was never found and eventually abandoned. Further scientific attention slowly began in the 1930s referring to slime as a microbial product and then was inspired by “How bacteria stick” by Costerton et al. in 1978, and the matrix material was considered to be polysaccharides. Later, it turned out that proteins, nucleic acids and lipids were major other constituents of the extracellular polymeric substances (EPS), an acronym which was highly discussed. The role of the EPS matrix turns out to be fundamental for biofilms, in terms of keeping cells in proximity and allowing for extended interaction, resource capture, mechanical strength and other properties, which emerge from the life of biofilm organisms, including enhanced tolerance to antimicrobials and other stress. The EPS components are extremely complex and dynamic and fulfil many functional roles, turning biofilms into the most ubiquitous and successful form of life on Earth.

Overview

The claims and statistics of this paper haven’t been checked yet.

Run the check — free, no account needed. It reads the open-access text and checks the sources it cites.

A few free checks a day · papers already verified open instantly · results are public

Integrity signals

Registry

Crossref · 10.3390/microorganisms4040041

Cites

96 of 96 references verified0 citation errors · 0 not checkable

Statistics

Figures

Figure 1 Bathybius Haeckelii. Figure 1 on Table 17 in Ernst Haeckels Article, Beiträge zur Plastidentheorie (1870) (Image in public domain).
Figure 1 Bathybius Haeckelii. Figure 1 on Table 17 in Ernst Haeckels Article, Beiträge zur Plastidentheorie (1870) (Image in public domain).CC BY · PubMed Central
Figure 2 Resource caption and retention by and in biofilms. The biofilm is a sponge-like system that provides surfaces for the sorption of a diverse range of molecules that can be sequestered from the environment. This confers several benefits to the biofilm, such as nutrient acquisition and matrix stabilization. Similarly, the physicochemical properties of the matrix enable biofilms to retain and stabilize extracellular digestive enzymes produced by biofilm cells, turning the matrix into an external digestive system. Surface-attached biofilms are not only able to take up nutrients from the water phase but can also digest biodegradable components from the substratum, which is exposed to enzymes in the matrix (after [19], with permission).
Figure 2 Resource caption and retention by and in biofilms. The biofilm is a sponge-like system that provides surfaces for the sorption of a diverse range of molecules that can be sequestered from the environment. This confers several benefits to the biofilm, such as nutrient acquisition and matrix stabilization. Similarly, the physicochemical properties of the matrix enable biofilms to retain and stabilize extracellular digestive enzymes produced by biofilm cells, turning the matrix into an external digestive system. Surface-attached biofilms are not only able to take up nutrients from the water phase but can also digest biodegradable components from the substratum, which is exposed to enzymes in the matrix (after [19], with permission).CC BY · PubMed Central
Figure 3 Fluorescent staining of a biofilm. Colour allocation: Nucleic acid stain (SybrGreen) = green; lectin stain (AAL-Alexa568) = red; autofluorescence of algae (chlorophyll A) = blue; autofluoresence of cyanobacteria = purple/white. Image dimensions: 246 × 246 μm (from [34], with permission, source: [35]).
Figure 3 Fluorescent staining of a biofilm. Colour allocation: Nucleic acid stain (SybrGreen) = green; lectin stain (AAL-Alexa568) = red; autofluorescence of algae (chlorophyll A) = blue; autofluoresence of cyanobacteria = purple/white. Image dimensions: 246 × 246 μm (from [34], with permission, source: [35]).CC BY · PubMed Central
Figure 4 Properties of biofilms emerging from life in the EPS matrix (after [19], with permission).
Figure 4 Properties of biofilms emerging from life in the EPS matrix (after [19], with permission).CC BY · PubMed Central
Figure 5 Control of biofilm formation by c-di-Guanidinemonophosphate (c-di-GMP). Concentration of c-di-GMP is regulated by DGC (Diguanylate cyclase; GGDEF domain: diguanylate cyclase), PDE 1 (Phosphodiesterase; EAL domain: diguanylate phosphodiesterase, linearizes c-di-GMP to 5′-pGpG), PDE (nonspecific cellular PDEs, further degrading 5′-pGpG to GMP), and PDE 2 (Phosphodiesterase; HD-GYP domain, metal dependent, unrelated to the EAL domain, linearizes c-di-GMP to 5′-pGpG, degrades 5′-pGpG further to GMP). pGpG—5′ phosphoguanylyl(3′,5′)guanosine; GMP—Guanosine-5-phosphate (from Krauss, G.-J., Nies, D. (Eds.): Ecological Biochemistry-Environmental and Interspecies Interactions today, VCH Weinheim, 2014, with permission).
Figure 5 Control of biofilm formation by c-di-Guanidinemonophosphate (c-di-GMP). Concentration of c-di-GMP is regulated by DGC (Diguanylate cyclase; GGDEF domain: diguanylate cyclase), PDE 1 (Phosphodiesterase; EAL domain: diguanylate phosphodiesterase, linearizes c-di-GMP to 5′-pGpG), PDE (nonspecific cellular PDEs, further degrading 5′-pGpG to GMP), and PDE 2 (Phosphodiesterase; HD-GYP domain, metal dependent, unrelated to the EAL domain, linearizes c-di-GMP to 5′-pGpG, degrades 5′-pGpG further to GMP). pGpG—5′ phosphoguanylyl(3′,5′)guanosine; GMP—Guanosine-5-phosphate (from Krauss, G.-J., Nies, D. (Eds.): Ecological Biochemistry-Environmental and Interspecies Interactions today, VCH Weinheim, 2014, with permission).CC BY · PubMed Central
Figure 6 The dammned polymer matrix (J.C. Bryers, Univ. Washington, with permission).
Figure 6 The dammned polymer matrix (J.C. Bryers, Univ. Washington, with permission).CC BY · PubMed Central

Verification

Claims the paper makes

The claims and statistics of this paper haven’t been checked yet.

Run the check — free, no account needed. It reads the open-access text and checks the sources it cites.

A few free checks a day · papers already verified open instantly · results are public

Claims other papers make about this one

Each is checked against this paper’s own text: does it say what the citing paper says it does?

Self-check

Percentages, sums and reported differences re-computed from the paper’s own numbers.

The self-check needs the paper’s full text.

Citations

Cites

Research works printed but not linked: 10

  1. Geesey, 1982, Microbial exopolymers: Ecological and economic considerations, ASM News, 48, 9
  2. Characklis, W.G., and Wilderer, P.A. (1989). Structure and Function of Biofilms, Wiley-Blackwell.
  3. Wingender, J., Neu, T.R., and Flemming, H.C. (1999). Microbial Extracellular Polymeric Substances, Springer. · DOI
  4. Wingender, J., Neu, T., and Flemming, H.-C. (1999). Microbial Extracellular Polymeric Substances, Springer. · DOI
  5. Flemming, H.-C., Neu, T.R., and Wingender, J. (2016). The Perfect Slime: Microbial Extracellular Polymeric Substances (EPS), IWA publishing. · DOI
  6. Moran, A., Brenan, P., Holst, O., and von Itzstein, M. (2009). Microbial Glycobiology: Structures, Relevance and Applications, Elsevier.
  7. Flemming, H.-C., Wingender, J., and Neu, T.R. (2016). The Perfect Slime—Microbial Extracellular Substances, IWA Publishing. · DOI
  8. Flemming, H.-C., Wingender, J., and Neu, T. (2016). The Perfect Slime—Microbial Extracellular Polymeric Substances, IWA Publishing. · DOI
  9. Flemming, H.-C., Neu, T.R., and Wingender, J. (2016). The Perfect Slime: Microbial Extracellular Polymeric Substances, Springer. · DOI
  10. Flemming, H.-C., Wingender, J., and Neu, T.R. (2016). The Perfect Slime—Microbial Extracellular Substances, IWA Publishing. · DOI

Cited by

Sorted by citations

See all 374

Metadata

Authors
Flemming, H.-C.
Affiliations
Nanyang Technological University (SG)
Corresponding author
Flemming, Hans-Curt
Venue
Microorganisms
Publisher
MDPI AG
Published
2016-11-18
Volume
4
Issue
4
Pages
41
ISSN
2076-2607 (ISSN-L)
OpenAlex
W2557171271
Cited by
374
References
96
Citations by year
2026: 31 · 2025: 49 · 2024: 55 · 2023: 38 · 2022: 50 · 2021: 35 · 2020: 52 · 2019: 32 · 2018: 20 · 2017: 12
FWCI
1.93
Type
journal article
Language
en
Open access
Locations
  • Microorganisms (publishedVersion, cc-by)
  • PubMed Central (publishedVersion, cc-by)
  • PubMed (publishedVersion)
  • DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion, cc-by-sa)
Topics
  • Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Bacterial biofilms and quorum sensing
  • Physical Sciences › Environmental Science › Pollution › Wastewater Treatment and Nitrogen Removal
  • Life Sciences › Agricultural and Biological Sciences › Food Science › Polysaccharides Composition and Applications
Keywords
extracellular polymeric substances, slime, biofilm, biofilm matrix, polysaccharides, extracellular proteins, extracellular nucleic acids, extracellular lipids, antimicrobial tolerance, Ernst Haeckel

About these checks

Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.

Statistics and arithmetic are recomputed, not judged.

Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.

What counts as a claim ›How we measure confidence ›How we check this ›

Are you an author, or do you see a problem with this page? Request a review