Earth's Systems / Biological processes / Terrestrial ecosystem functioning / Living soil system / Saprotrophic fungi / White-rot and brown-rot fungi: two routes to lignin and cellulose decay

White-rot and brown-rot fungi: two routes to lignin and cellulose decay

How wood and litter fungi attack lignified plant material. White-rot fungi use peroxidases and laccases to oxidize lignin; brown-rot fungi use Fenton chemistry to depolymerize cellulose and leave modified lignin. Mineral nitrogen from decomposition feeds back by repressing ligninolytic enzymes.

Boundary: where this process starts

Starts with: White-rot basidiomycetes secrete class II peroxidases (lignin, manganese, versatile) and laccases that oxidize lignin and leave a pale, cellulose-rich residue

Boundary: where it ends

Ends after: Hypothesis: class II peroxidases arose about 295 million years ago, coinciding with the end of major coal formation (competing explanations exist); Partly oxidized lignin fragments and fungal melanin enter the dissolved pool and can become humic precursors

PARTICIPANTSRESOURCESWhite-rot fungiBrown-rot fungiSaprotrophicfungiLigninperoxidaseLaccaseOtherparticipantsLigninHydrogenperoxideCelluloseManganeseOxalateImpact +3 of 5. Click for details and evidence.White-rot routeImpact +3 of 5. Click for details and evidence.Brown-rot routeImpact +3 of 5. Click for details and evidence.Mn peroxidase adds a diffusiImpact +2 of 5. Click for details and evidence.Lignin removal is co-metabolImpact 01 of 5. Click for details and evidence.Evolutionary originImpact +3 of 5. Click for details and evidence.Oxidized fragments releasedImpact +3 of 5. Click for details and evidence.Modified lignin residue and Impact +2 of 5. Click for details and evidence.Lignin fragments accumulateImpact +2 of 5. Click for details and evidence.Decomposition releases NImpact -3 of 5. Click for details and evidence.↻ balances · moderate · 0.25 High mineral N Impact 01 of 5. Click for details and evidence.Context for the modern carboImpact +4 of 5. Click for details and evidence.Carbon released as CO2 and dWhite-rot basidiomycetes secrete class II peroxidases (lignin, manganese, versatile) and laccases that oxidize lignin and leave a pale, cellulose-rich residue: uses up Ligninuses upWhite-rot basidiomycetes secrete class II peroxidases (lignin, manganese, versatile) and laccases that oxidize lignin and leave a pale, cellulose-rich residue: uses up Hydrogen peroxideuses upManganese peroxidase oxidizes Mn(II) to Mn(III) chelated by oxalate, a diffusible oxidant that attacks lignin beyond the enzyme's reach: uses up Ligninuses upManganese peroxidase oxidizes Mn(II) to Mn(III) chelated by oxalate, a diffusible oxidant that attacks lignin beyond the enzyme's reach: uses up Manganeseuses upManganese peroxidase oxidizes Mn(II) to Mn(III) chelated by oxalate, a diffusible oxidant that attacks lignin beyond the enzyme's reach: uses up Oxalateuses upBrown-rot fungi lack class II peroxidases and use Fenton chemistry (Fe2+ + H2O2 gives hydroxyl radicals) to depolymerize cellulose, leaving modified lignin: uses up Ligninuses upBrown-rot fungi lack class II peroxidases and use Fenton chemistry (Fe2+ + H2O2 gives hydroxyl radicals) to depolymerize cellulose, leaving modified lignin: uses up Hydrogen peroxideuses upBrown-rot fungi lack class II peroxidases and use Fenton chemistry (Fe2+ + H2O2 gives hydroxyl radicals) to depolymerize cellulose, leaving modified lignin: uses up Celluloseuses upHypothesis: class II peroxidases arose about 295 million years ago, coinciding with the end of major coal formation (competing explanations exist): uses LigninLignin is co-metabolized: fungi need cellulose or another carbon source and H2O2 from oxidases; lignin alone does not support growth: uses up Ligninuses upLignin is co-metabolized: fungi need cellulose or another carbon source and H2O2 from oxidases; lignin alone does not support growth: uses up Hydrogen peroxideuses upLignin is co-metabolized: fungi need cellulose or another carbon source and H2O2 from oxidases; lignin alone does not support growth: uses up Celluloseuses upPartly oxidized lignin fragments and fungal melanin enter the dissolved pool and can become humic precursors: uses up Ligninuses upMineral nitrogen released as litter decays represses ligninolytic enzymes and phenol oxidase in lignin-rich litter, slowing late-stage decay while speeding early cellulose decay: uses Ligninoxidized by peroxidasesand laccasesLignin: oxidized by peroxidases and laccasesconsumed by peroxidasereactionsHydrogen peroxide: consumed by peroxidase reactionsattacked by diffusibleoxidantLignin: attacked by diffusible oxidantoxidized to Mn(III) byenzymeManganese: oxidized to Mn(III) by enzymechelates Mn(III) asoxidantOxalate: chelates Mn(III) as oxidantmodified rather thanremovedLignin: modified rather than removedreacts with iron to formradicalsHydrogen peroxide: reacts with iron to form radicalsdepolymerized byhydroxyl radicalsCellulose: depolymerized by hydroxyl radicalsoxidized by evolvedperoxidasesLignin: oxidized by evolved peroxidasesco-metabolized alongsidecelluloseLignin: co-metabolized alongside cellulosesupplied by oxidasesHydrogen peroxide: supplied by oxidasessupplies carbon forgrowthCellulose: supplies carbon for growthfragmented into humicprecursorsLignin: fragmented into humic precursorsdecay slowed by nitrogenLignin: decay slowed by nitrogenStart: White-rot andbrown-rot fungi: tworoutes to lignin a…STARTStart: White-rot and brown-rot fungi: two routes to lignin and cellulose decayEnd: White-rot andbrown-rot fungi: tworoutes to lignin a…ENDEnd: White-rot and brown-rot fungi: two routes to lignin and cellulose decayWhite-rotbasidiomycetes secreteclass II peroxidas…📄1White-rot basidiomycetes secrete class II peroxidases (lignin, manganese, versatile) and laccases that oxidize lignin and leave a pale, cellulose-rich residueManganese peroxidaseoxidizes Mn(II) toMn(III) chelated …📄1Manganese peroxidase oxidizes Mn(II) to Mn(III) chelated by oxalate, a diffusible oxidant that attacks lignin beyond the enzyme's reachBrown-rot fungi lackclass II peroxidasesand use Fent…📄1Brown-rot fungi lack class II peroxidases and use Fenton chemistry (Fe2+ + H2O2 gives hydroxyl radicals) to depolymerize cellulose, leaving modified ligninHypothesis: class IIperoxidases arose about295 million years ag…📄1Hypothesis: class II peroxidases arose about 295 million years ago, coinciding with the end of major coal formation (competing explanations exist)Lignin isco-metabolized: fungineed cellulose …📄1Lignin is co-metabolized: fungi need cellulose or another carbon source and H2O2 from oxidases; lignin alone does not support growthPartly oxidized ligninfragments and fungalmelanin enter t…📄1Partly oxidized lignin fragments and fungal melanin enter the dissolved pool and can become humic precursorsMineral nitrogenreleased as litterdecays repress…📄1Mineral nitrogen released as litter decays represses ligninolytic enzymes and phenol oxidase in lignin-rich litter, slowing late-stage decay while speeding early cellulose decay
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)
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White-rot basidiomycetes secrete class II peroxidases (lignin, manganese, versatile) and laccases that oxidize lignin and leave a pale, cellulose-rich residue
Evidence (1) · added by Ember
  • Floudas et al. 2012, Science 336:1715: Class II peroxidases are the signature enzymes of white-rot Agaricomycetes. (confidence 0.85)
Manganese peroxidase oxidizes Mn(II) to Mn(III) chelated by oxalate, a diffusible oxidant that attacks lignin beyond the enzyme's reach
Evidence (1) · added by Ember
  • Hofrichter 2002, Enzyme and Microbial Technology 30:454, Review: lignin conversion by manganese peroxidase: Chelated Mn(III) acts as a diffusible mediator oxidizing phenolic lignin units. (confidence 0.85)
Brown-rot fungi lack class II peroxidases and use Fenton chemistry (Fe2+ + H2O2 gives hydroxyl radicals) to depolymerize cellulose, leaving modified lignin
Evidence (1) · added by Ember
  • Eastwood et al. 2011, Science 333:762, The plant cell wall-decomposing machinery underlies the functional diversity of forest fungi: Brown-rot genomes have reduced oxidative and cellulolytic enzyme sets and rely on oxidative radical chemistry. (confidence 0.85)
Hypothesis: class II peroxidases arose about 295 million years ago, coinciding with the end of major coal formation (competing explanations exist)
Evidence (1) · added by Ember
  • Floudas et al. 2012, Science 336:1715: Molecular clock places the origin of lignin-degrading peroxidases near the Carboniferous-Permian boundary. (confidence 0.6)
Lignin is co-metabolized: fungi need cellulose or another carbon source and H2O2 from oxidases; lignin alone does not support growth
Evidence (1) · added by Ember
  • Kirk & Farrell 1987, Annual Review of Microbiology 41:465, Enzymatic combustion: the microbial degradation of lignin: Ligninolysis is a secondary-metabolic, co-metabolic process that exposes cellulose rather than feeding on lignin. (confidence 0.8)
Partly oxidized lignin fragments and fungal melanin enter the dissolved pool and can become humic precursors
Evidence (1) · added by Ember
  • Kogel-Knabner 2002, Soil Biology and Biochemistry 34:139, The macromolecular organic composition of plant and microbial residues as inputs to soil organic matter: Lignin-derived and microbial residues contribute aromatic and phenolic structures to soil organic matter. (confidence 0.7)
Mineral nitrogen released as litter decays represses ligninolytic enzymes and phenol oxidase in lignin-rich litter, slowing late-stage decay while speeding early cellulose decay
Evidence (1) · added by Ember
  • Carreiro, Sinsabaugh, Repert & Parkhurst 2000, Ecology 81:2359, Microbial enzyme shifts explain litter decay responses to simulated nitrogen deposition: N additions raised cellulase but lowered phenol oxidase activity in high-lignin litter, slowing late-stage decay. (confidence 0.8)