Biofilms and quorum signalling in soil microcolonies

How soil bacteria attach to particles and roots, embed themselves in an extracellular polymer matrix, and use diffusible signals such as acyl-homoserine lactones to coordinate gene expression when signal accumulates. It covers diffusion limits on signalling, signal degradation, cheaters, and the effects on desiccation tolerance, aggregation and gene transfer.

Boundary: where this process starts

Starts with: Bacterial cells cover under 1% of mineral surface area, so most cells are isolated and microcolonies are patchy

Boundary: where it ends

Ends after: EPS and cells glue mineral particles into aggregates and raise aggregate stability; Close packing and the matrix raise gene transfer: conjugative plasmids can induce biofilm formation

PARTICIPANTSRESOURCESSoil bacteriaBiofilmPseudomonasSoil waterMineralsurfacesOtherparticipantsCarbonAcyl-homoserinelactonesExtracellularpolysaccharidesWaterPore spaceImpact 02 of 5. Click for details and evidence.patchy colonisationImpact 02 of 5. Click for details and evidence.soil pH and temperatureImpact 02 of 5. Click for details and evidence.quenching communityImpact 02 of 5. Click for details and evidence.dry-wet cyclesImpact +3 of 5. Click for details and evidence.surface contactImpact +3 of 5. Click for details and evidence.microcolony growsImpact +3 of 5. Click for details and evidence.local density and confinemenImpact +4 of 5. Click for details and evidence.35% signal accumulatesImpact -3 of 5. Click for details and evidence.55% dilutedImpact -3 of 5. Click for details and evidence.10% destroyedImpact +4 of 5. Click for details and evidence.AHL binds receptorImpact +4 of 5. Click for details and evidence.↻ reinforces · strong · 0.5 autoinduction ofImpact +4 of 5. Click for details and evidence.target genes onImpact -3 of 5. Click for details and evidence.lactone hydrolysisImpact -3 of 5. Click for details and evidence.↻ balances · moderate · 0.3 signal degradedImpact -3 of 5. Click for details and evidence.public goods attract cheaterImpact -3 of 5. Click for details and evidence.↻ balances · moderate · 0.3 public goods proImpact +3 of 5. Click for details and evidence.more EPSImpact +2 of 5. Click for details and evidence.dense communityImpact +2 of 5. Click for details and evidence.water-holding matrixImpact -2 of 5. Click for details and evidence.nutrients later declineImpact +2 of 5. Click for details and evidence.dispersed cells recoloniseImpact +2 of 5. Click for details and evidence.leaveImpact +3 of 5. Click for details and evidence.aggregate and desiccation beImpact 01 of 5. Click for details and evidence.gene transferImpact +2 of 5. Click for details and evidence.dispersedBacterial cells cover under 1% of mineral surface area, so most cells are isolated and microcolonies are patchy: uses Pore spaceCells attach to a surface using pili, flagella and adhesins, then secrete extracellular polymeric substances (EPS): uses CarbonCells attach to a surface using pili, flagella and adhesins, then secrete extracellular polymeric substances (EPS): makes Extracellular polysaccharidesmakesMicrocolony grows, matrix thickens, cell density in the pore rises: uses up Carbonuses upSignal fate in the pore: accumulates above threshold, is diluted away, or is destroyed: uses Acyl-homoserine lactonesSignal fate in the pore: accumulates above threshold, is diluted away, or is destroyed: uses WaterGram-negative cells make acyl-homoserine lactone (AHL) with LuxI-type synthases: uses up Carbonuses upGram-negative cells make acyl-homoserine lactone (AHL) with LuxI-type synthases: makes Acyl-homoserine lactonesmakesAHL binds a LuxR-type receptor above a threshold and the complex switches on target genes, including luxI itself (autoinduction): uses Acyl-homoserine lactonesAlkaline pH and warmth hydrolyse the AHL lactone ring, so signals persist longer in acid, cool soil: uses up Acyl-homoserine lactonesuses upQuorum quenching: AHL lactonases (AiiA) and plant signal mimics degrade or confuse the signal: uses up Acyl-homoserine lactonesuses upSignal-blind cheaters use public goods without paying the cost, and may spread: uses CarbonEPS retains water and buffers drying, so matrix-rich cells survive desiccation better: uses Extracellular polysaccharidesEPS retains water and buffers drying, so matrix-rich cells survive desiccation better: uses WaterEPS and cells glue mineral particles into aggregates and raise aggregate stability: uses Extracellular polysaccharidesNutrient decline signals cells to leave (lower cyclic-di-GMP) and disperse to new surfaces: uses CarbonSignal diluted below threshold: cells remain solitary or in loose microcolonies: uses Watercolonised only inpatchesPore space: colonised only in patchesinvested in surfaceattachmentCarbon: invested in surface attachmentsecreted as matrixExtracellular polysaccharides: secreted as matrixaccumulates inmicrocoloniesCarbon: accumulates in microcoloniesaccumulate or degrade inporeAcyl-homoserine lactones: accumulate or degrade in poredilutes or carriessignal awayWater: dilutes or carries signal awayinvested in signalproductionCarbon: invested in signal productionsynthesised by LuxIenzymesAcyl-homoserine lactones: synthesised by LuxI enzymesbinds receptor abovethresholdAcyl-homoserine lactones: binds receptor above thresholdhydrolysed at alkalinepHAcyl-homoserine lactones: hydrolysed at alkaline pHdegraded by lactonasesAcyl-homoserine lactones: degraded by lactonasesused without paying costCarbon: used without paying costretain water duringdryingExtracellular polysaccharides: retain water during dryingretained in matrixWater: retained in matrixglue particles intoaggregatesExtracellular polysaccharides: glue particles into aggregatessignals cells todisperseCarbon: signals cells to dispersesignal diluted belowthresholdWater: signal diluted below thresholdStart: Cells land on aparticle or rootsurface in a water filmSTARTStart: Cells land on a particle or root surface in a water filmEnd: Collectivebehaviour expressed orcells dispersedENDEnd: Collective behaviour expressed or cells dispersedBacterial cells coverunder 1% of mineralsurface area, so mo…📄1Bacterial cells cover under 1% of mineral surface area, so most cells are isolated and microcolonies are patchyCells attach to asurface using pili,flagella and adhesin…📄1Cells attach to a surface using pili, flagella and adhesins, then secrete extracellular polymeric substances (EPS)Microcolony grows,matrix thickens, celldensity in the po…📄1Microcolony grows, matrix thickens, cell density in the pore risesSignal fate in thepore: accumulates abovethreshold, is dilut…📄1Signal fate in the pore: accumulates above threshold, is diluted away, or is destroyedGram-negative cellsmake acyl-homoserinelactone (AHL) wi…📄1Gram-negative cells make acyl-homoserine lactone (AHL) with LuxI-type synthasesAHL binds a LuxR-typereceptor above athreshold and t…📄1AHL binds a LuxR-type receptor above a threshold and the complex switches on target genes, including luxI itself (autoinduction)Quorum-controlledoutputs: antibiotics(phenazines…📄1Quorum-controlled outputs: antibiotics (phenazines), exoenzymes, siderophores, more EPSAlkaline pH and warmthhydrolyse the AHLlactone ring, …📄1Alkaline pH and warmth hydrolyse the AHL lactone ring, so signals persist longer in acid, cool soilQuorum quenching: AHLlactonases (AiiA) andplant signal mimi…📄1Quorum quenching: AHL lactonases (AiiA) and plant signal mimics degrade or confuse the signalSignal-blind cheatersuse public goodswithout paying t…📄1Signal-blind cheaters use public goods without paying the cost, and may spreadEPS retains water andbuffers drying, somatrix-rich cel…📄1EPS retains water and buffers drying, so matrix-rich cells survive desiccation betterEPS and cells gluemineral particles intoaggregates and rai…📄1EPS and cells glue mineral particles into aggregates and raise aggregate stabilityClose packing and thematrix raise genetransfer: conjugati…📄1Close packing and the matrix raise gene transfer: conjugative plasmids can induce biofilm formationNutrient declinesignals cells to leave(lower cyclic-di-GM…📄1Nutrient decline signals cells to leave (lower cyclic-di-GMP) and disperse to new surfacesSignal diluted belowthreshold: cells remainsolitary or in loo…📄1Signal diluted below threshold: cells remain solitary or in loose microcolonies
Line thickness = size of impact (1 to 5)% on a line = chance a fork takes that branch📄 = pieces of evidence on a stepNegative effectLoops backHypothesisResource used up (arrow into the step)Resource used, not used upResource made or released (arrow out of the step)
What am I looking at? The concepts on every map
Participant
Someone or something that takes part in a process or influences it: a plant, a microbe, a river, a machine, a force. Participants are the rows of a map.
Resource
Something a step uses or uses up: water, oxygen, sunlight, a tool. Each step has an arrow down to the resources it uses.
Step
One thing that happens in a process. Every step has at least one participant and carries evidence.
Decision fork
A step where the process can go several ways. Every outcome is labeled with a percentage, and the percentages add up to 100%. It is not limited to yes or no.
Loop
A line that carries the process back to an earlier step. We record whether it reinforces or balances, how strong it is (weak, moderate, strong), and its gain (0 to 1): the share of the effect that returns on each pass.
Size of impact
Every line says how much the earlier step affects the next one (1 to 5), in which direction (+, -, 0), and why. Thicker lines mean bigger impact.
Evidence
The support for a claim: a source and what it shows. Every step, line, change and vote needs it. Click any step or line to read its evidence.
Boundary
Where a process starts and where it ends. Every process, and every subprocess, has a start and an end.
Subprocess
A step that is itself a whole process, with its own boundary, steps and lines. Click a step marked ↘ to open it. Subprocesses nest as deep as the evidence goes.
Description
A plain-language summary of what a process is and what it covers.
Hypothesis
A proposed new connection between two parts of the map that nobody has linked. It carries a prediction, a way to test it, and a count of papers on each topic and on both. It is an unverified idea until someone tests it.

Every step, with its evidence

The same information as the map, as a list you can search and share. Click a participant or resource to see everywhere it appears.

Bacterial cells cover under 1% of mineral surface area, so most cells are isolated and microcolonies are patchy
Evidence (1) · added by Ember-C
  • Young, Crawford, Nunan, Otten & Spiers 2008, Advances in Agronomy 100:81-121 (Microbial distribution in soils: physics and scaling): Microbial cells occupy a very small fraction of the available soil surface and are clustered in microcolonies around organic substrates. (confidence 0.55)
Cells attach to a surface using pili, flagella and adhesins, then secrete extracellular polymeric substances (EPS)
Evidence (1) · added by Ember-C
  • Flemming & Wingender 2010, Nature Reviews Microbiology 8:623-633: Attachment and EPS production are early steps in biofilm formation; EPS can be 50-90% of the biofilm organic carbon. (confidence 0.8)
Microcolony grows, matrix thickens, cell density in the pore rises
Evidence (1) · added by Ember-C
  • Flemming & Wingender 2010, Nature Reviews Microbiology 8:623-633: Biofilms grow as dense multicellular aggregates in a self-produced matrix. (confidence 0.8)
⑂ Signal fate in the pore: accumulates above threshold, is diluted away, or is destroyed
Evidence (1) · added by Ember-C
  • Redfield 2002, Trends in Microbiology 10:365-370 (Is quorum sensing a side effect of diffusion sensing?): Whether a diffusible signal accumulates depends on diffusion and mass transfer as well as cell density, so quorum may be a measure of confinement. Shares are an estimate for a typical soil colony and are not measured. (confidence 0.4)
Gram-negative cells make acyl-homoserine lactone (AHL) with LuxI-type synthases
Evidence (1) · added by Ember-C
  • Fuqua, Winans & Greenberg 1994, Journal of Bacteriology 176:269-275 (Quorum sensing in bacteria: the LuxR-LuxI family of cell density-responsive transcriptional regulators): LuxI-type enzymes synthesise AHL signals that accumulate with cell density. (confidence 0.9)
AHL binds a LuxR-type receptor above a threshold and the complex switches on target genes, including luxI itself (autoinduction)
Evidence (1) · added by Ember-C
  • Fuqua, Winans & Greenberg 1994, Journal of Bacteriology 176:269-275: LuxR-AHL activates transcription including the signal synthase gene, creating positive feedback. (confidence 0.9)
Quorum-controlled outputs: antibiotics (phenazines), exoenzymes, siderophores, more EPS
Evidence (1) · added by Ember-C
  • Pierson, Wood & Pierson 1998, Annual Review of Phytopathology 36:207-225 (Homoserine lactone-mediated gene regulation in plant-associated bacteria): Phenazine antibiotic production by Pseudomonas aureofaciens on wheat roots is regulated by AHL signalling. (confidence 0.8)
Alkaline pH and warmth hydrolyse the AHL lactone ring, so signals persist longer in acid, cool soil
Evidence (1) · added by Ember-C
  • Yates et al. 2002, Infection and Immunity 70:5635-5646 (N-acylhomoserine lactones undergo lactonolysis in a pH-, temperature-, and acyl chain length-dependent manner): AHL half-life drops sharply as pH rises above neutral and with higher temperature and shorter acyl chains. (confidence 0.85)
Quorum quenching: AHL lactonases (AiiA) and plant signal mimics degrade or confuse the signal
Evidence (1) · added by Ember-C
  • Dong et al. 2000, PNAS 97:3526-3531 (AiiA, an enzyme that inactivates the acylhomoserine lactone quorum-sensing signal): The AiiA lactonase from Bacillus inactivates AHL signals by hydrolysis. (confidence 0.85)
Signal-blind cheaters use public goods without paying the cost, and may spread
Evidence (1) · added by Ember-C
  • Diggle, Griffin, Campbell & West 2007, Nature 450:411-414 (Cooperation and conflict in quorum-sensing bacterial populations): Quorum-sensing mutants can exploit cooperative production of public goods and reduce productivity, unless cooperation is stabilised. (confidence 0.85)
EPS retains water and buffers drying, so matrix-rich cells survive desiccation better
Evidence (1) · added by Ember-C
  • Roberson & Firestone 1992, Applied and Environmental Microbiology 58:1284-1291 (Relationship between desiccation and exopolysaccharide production in a soil Pseudomonas sp.): EPS-rich Pseudomonas cells held more water and survived desiccation better than low-EPS cells. (confidence 0.85)
EPS and cells glue mineral particles into aggregates and raise aggregate stability
Evidence (1) · added by Ember-C
  • Costa, Raaijmakers & Kuramae 2018, Frontiers in Microbiology 9:1636 (Microbial extracellular polymeric substances: ecological function and impact on soil aggregation): EPS bind particles and contribute to soil aggregation and water retention. (confidence 0.8)
Close packing and the matrix raise gene transfer: conjugative plasmids can induce biofilm formation
Evidence (1) · added by Ember-C
  • Ghigo 2001, Nature 412:442-445 (Natural conjugative plasmids induce bacterial biofilm development): Conjugative plasmids encode factors that promote biofilm formation, favouring their own transfer. (confidence 0.8)
Nutrient decline signals cells to leave (lower cyclic-di-GMP) and disperse to new surfaces
Evidence (1) · added by Ember-C
  • McDougald et al. 2012, Nature Reviews Microbiology 10:39-50 (Should we stay or should we go: mechanisms and ecological consequences for biofilm dispersal): Starvation and other cues trigger biofilm dispersal through signalling that lowers cyclic-di-GMP. (confidence 0.8)
Signal diluted below threshold: cells remain solitary or in loose microcolonies
Evidence (1) · added by Ember-C
  • Redfield 2002, Trends in Microbiology 10:365-370: In open, well-mixed water films signal does not build up, so the response is not triggered. (confidence 0.5)