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A list of cellulolytic bacteria Page

This list contains all truly cellulolytic bacterial species described in the literature (I am aware of
hydrolyzing crystalline cellulose and/or fermenting crystalline cellulose as sole carbon source.
hydrolysis means continuous hydrolysis of at least microcrystalline cellulose like Avicel, or bet
linters, bacterial cellulose or the like - this is more than just e.g. initial activity on Avicel (e.g. rel
equivalents).

The species are arranged by a superposition of the phylogenetic tree derived from calculation o
sequences with the ARB program package (go to the Microbiology homepage), and the new taxono
in Bergeys Manual of Systematic Bacteriology (Garrity, 2001).

Such a list will never be complete. So if you have data on a bacterial species which clearly show
crystalline cellulose, please let me know.
Wolfgang H. Schwarz (e-mail)

Phylogeny

Genus

Species

Temp

Source

hot spring

Phylum Firmicutes, Class Clostridia, Order Clostridiales


Fam.
Syntrophomonodaceae

Caldocellulosiruptor

saccharolyticus

Caldocellulosiruptor

lactoaceticus

Caldocellulosiruptor

kristjanssonii

"Anaerocellum"

thermophilum

Butyrivibrio

fibrisolvens

rumen

Ruminococcus

flavefaciens

rumen

Ruminococcus

succinogenes

rumen

Ruminococcus

albus

rumen

Fam. Eubacteriaceae

Eubacterium

cellulolyticum

rumen

Fam. Clostridiaceae

Clostridium

acetobutylicum

soil

Fam. Lachnospiraceae

hot spring

Clostridium

chartatabidum

rumen

Clostridium

cellulovorans

wood
fermenter

Clostridium

herbivorans

pig
intestine

Clostridium

cellulosi

manure

Clostridium

cellobioparum

rumen

Clostridium

papyrosolvens

paper mill

Clostridium

josui

compost

Clostridium

cellulolyticum

compost

Clostridium

aldrichii

wood
fermenter

Clostridium

stercorarium

compost

Clostridium

thermocellum

sewage +
soil

Clostridium

cellulofermentans

manure

Clostridium

celerescens

manure

Clostridium

thermopapyrolyticum

mud

Clostridium

thermocopriae

hot spring

Clostridium

sp. C7

mud

Bacteroides

sp. P-1

rotting
biomass

Bacteroides

cellulosolvens

sewage

Acetivibrio

cellulolyticus

sewage

Acetivibrio

cellulosolvens

sewage

Phylum Firmicutes, Class Bacilli, Order Bacillales


Fam.
Thermoactinomyces
Thermoactinomycetaceae

Fam. Bacillaceae

sp. YX

Caldibacillus

cellulovorans

Bacillus

circulans

Phylum Actinobacteria, Order Actinomycetales


Acidothermus

cellulolyticus

Cellulomonas

biazotea

Cellulomonas

cartae

Cellulomonas

cellasea

Cellulomonas

cellulans

soil

Cellulomonas

fimi

soil

Cellulomonas

flavigena

soil

Cellulomonas

gelida

Cellulomonas

iranensis

forest soil

Cellulomonas

persica

forest soil

Cellulomonas

uda

sewage

Subord. Frankineae,

acidic hot
spring

Fam.
Acidothermaceae

Subord.
Micromonosporineae,

Fam.
Cellulomonadaceae

Curtobacterium

falcumfaciens

soil

Fam. Microbacteriaceae

Micromonospora

melonosporea

compost

Fam.
Micromonosporaceae

Actinoplanes

aurantiaca

soil

Streptomyces

reticuli

soil

Streptomyces

alboguseolus

Suborder
Streptomycineae,

Streptomyces

aureofaciens

Fam.
Streptomycetaceae

Streptomyces

cellulolyticus

Streptomyces

flavogriseus

Streptomyces

lividans

Streptomyces

nitrosporeus

Streptomyces

olivochromogenes

Streptomyces

rochei

Streptomyces

thermovulgaris

Streptomyces

viridosporus

Thermobifida

alba

Thermobifida(Thermomonospora)

fusca

Thermobifida

cellulolytica

Thermomonospora

curvata

Microbispora

bispora

compost

soil

termite
gut

Subord.
Streptosporangiaceae,

Fam.
Nocardiopsaceae

h
soil

compost

soil

Fam.
Streptosporangiaceae

Phylum Fibrobacteres, Class Fibrobacteres, Order Fibrobacterales


Fam. Fibrobacteriaceae

Fibrobacter

succinogenes

rumen

Phylum Bacteroidetes, Class Sphingobacteria, Order Sphingobacteriales


Fam. Flexibacteriaceae

Sporocytophaga

myxococcoides

Cytophaga

sp.

m
m

soil
soil

Phylum Bacteroidetes, Class Flavobacteria, Order Flavobacteriales


Fam. Flavobacteriaceae

Flavobacterium

johnsoniae

soil

Phylum Proteobacteria, Class Betaproteobacteria, Order Burkholderiales


Fam. Alcaligenaceae

Achromobacter

piechaudii

soil

Phylum Proteobacteria, Class Gammaproteobacteria, Order Xanthomonadales


Fam. Xanthomonadaceae

Xanthomonas

sp.

brack
water

Phylum Proteobacteria, Class Gammaproteobacteria, Order Pseudomonadales


Fam. Pseudomonaceae

Cellvibrio

vulgaris, fulvus

soil

Cellvibrio

gilvus

soil

Cellvibrio

mixtus

soil

Pseudomonas

fluorescens
(cellulosa)

plant
pathogen

Pseudomonas

mendocina

soil

Phylum Proteobacteria, Class Deltaproteobacteria


Fam. unclassified

Myxobacter

sp. AL-1

soil

+d

Last update 24.04.2003, W. Schwarz, TUM, Microbiology

Legend: Temp, growth temperature: m, mesophilic; h, thermophilic (growth optimum above 50 C).
Co, presence of cellulosomes; evidence: a) presence of dockerin of cohesin sequences; b)
biochemical evidence (multienzyme complexes); c) reaction with Ct-CipA-scaffoldin antibodies; d)
presence of cell protuberances in electron microscopy.

Acknowledgements: I thank all collegues who contributed to this page


for personal communication of unpublished results.

References:

A number of reviews has been published recently giving an overview:

1. Bayer EA, Morag E, Lamed R, Yaron S, Shoham Y (1998) Cellulosome structure: fourpronged attack using biochemistry, molecular biology, crystallography and bioinformatics.
In Carbohydrates from Trichoderma reesei and other microorganisms. (M. Claeyssens,
W. Nerinckx and K. Piens, eds.). pp. 39-65. The Royal Society of Chemistry, London
2. Coughlan MP, Mayer F (1992) The cellulose-decomposing bacteria and their enzyme
systems. In The Prokaryotes: a handbook on the biology of bacteria. 2ns edn., p. 460516. Edited by Balows A, Trper HG, Dworkin M, Harder W, Schleifer KH. SpringerVerlag, New York
3. Doi RH, Goldstein M, Hashida S, Park JS, Takagi M (1994) The Clostridium
cellulovorans cellulosome. Crit Rev Microbiol 20: 87-93
4. Doi RH, Park JS, Liu CC, Malburg LM, Tamaru Y, Ichiishi A, Ibrahim A (1998) Cellulosome
and noncellulosomal cellulases of Clostridium cellulovorans. Extremophiles 2: 53-60
5. Gal L, Pags S, Gaudin C, Belaich A, Reverbel-leroy C, Tardif C, Belaich JP (1997)
Characterization of the cellulolytic complex (cellulosome) produced by Clostridium
cellulolyticum. Appl Environ Microbiol 63: 903-909
6. Guglielmi G, Bguin P (1998) Cellulase and hemicellulase genes of Clostridium
thermocellum from five independent collections contain few overlaps and are widely
scattered across the chromosome. FEMS Microbiol Let 161: 209-215
7. Himmel ME, Ruth MF, Wyman CE (1999) Cellulase for commodity products from
cellulosic biomass. Curr Opin Biotechnol 10: 358-364
8. Karita S, Sakka K, Ohmiya K (1997) Cellulosomes, cellulase complexes, of anaerobic
microbes: their structure, models and functions. In Rumen Microbes and Digestive
Physiology in Ruminants, Onodera R et al. eds., pp. 47-57. Japan Sci. Soc. Press,
Tokyo/S. Karger, Basel.

9. Schlein M (2000) Protein engineering of cellulases. Biochim Biophys Acta 1543: 239252
10. Sheehan J, Himmel M (1999) Enzymes, energy, and the environment: a strategic
perspective on the U.S. department of energys research and development activities for
bioethanol. Biotechnol Prog 15: 817-827
11. Shoham Y, Lamed R, Bayer EA (1999) The cellulosome concept as an efficient microbial
strategy for the polysaccharide degradation of insoluble polysaccharides. Trends
Microbiol 7: 275-281
12. Tomme P, Boraston A, McLean B, Kormos J, Creagh AL, Sturch K, Gilkes NR, Haynes
CA, Warren RAJ, Kilburn DG (1998) Characterization and affinity applications of
cellulose-binding domains. J Chromatogr B 715: 283-296
13. Tomme P, Warren RA, Miller RC Jr, Kilburn DG, Gilkes,N (1995) In: Enzymatic
Degradation of Insoluble Polysaccharides (Saddler, J.N. & Penner, M., eds.), Cellulosebinding domains: classification and properties. pp. 142-163, American Chemical Society,
Washington

A server containing all known cellulase sequences, grouped


in families. It is continuously updated:
14. Coutinho, P.M. & Henrissat, B. (1999) Carbohydrate-Active Enzymes server at
URL: http://afmb.cnrs-mrs.fr/~cazy/CAZY/index.html
15. Henrissat B, Teeri TT, Warren RAJ (1998) A scheme for designating enzymes that
hydrolyse the polysaccharides in the cell walls of plants. FEBS Let 425: 352-354
16. Henrissat B., Bairoch A. (1996) Updating the sequence-based classification of glycosyl
hydrolases. Biochem. J. 316: 695-696.

These references cite the paper where the cellulolytic trait was
described:
17. Anderson KL, Blair BG (1996) Regulation of the cellulolytic activity of Eubacterium
cellulosolvens 5494: a review. SAAS Bull Biochem Biotechnol 9: 57-62
18. Aurilia V, Martin JC, McCrae SI, Scot KP, Rincon MT, Flint HJ (2000) Three multidomain
esterases from the cellulolytic rumen anaerobe Ruminococcus flavefaciens 17 that carry
divergent dockerin sequences. Microbiol 146: 1391-1397
19. Avitia CI, Castellanos-Juarez FX, Sanchez E, Tellez-Valencia A, Fajardo-Cavazos P,
Nicholson WL, Pedraza-Reyes M (2000) Temporal secretion of a multicellulolytic system
in myxobacter sp. AL-1 molecular cloning and heterologous expression of cel9 encoding
a modular endocellulase clustered in an operon with cel48, an exocellobiohydrolase
gene. Eur J Biochem. 267:7058-64

20. Blaich JP, Tardif C, Blaich A, Gaudin C (1997) The cellulolytic system of Clostridium
cellulolyticum. J Biotechnol 57: 3-14
21. Berger E, Jones WA, Jones DT, Woods DR (1990) Sequencing and expression of a
cellodextrinase (ced1) gene from Butyrivibrio fibrisolvens H17c cloned in Escherichia coli.
Mol Gen Genet 223: 310-318
22. Blackall LL, Hayward AC, Sly LI (1985) Cellulolytic and extremophilic Gram-negative
bacteria: revival of the genus Cellvibrio. J Appl Bacteriol 59: 81-97
23. Bredholt S, Sonne-Hansen J, Nielsen P, Mathrani IM, Ahring BK
(1999) Caldicellulosiruptor kristjanssoniisp. nov., a cellulolytic, extremely thermophilic,
anaerobic bacterium. Int J Syst Bacteriol 49: 991-996
24. Cavedon K, Leschine SB, Canale-Parola E (1990) Cellulase system of a free-living,
mesophilic Clostridium(strain C7). J Bacteriol 172: 4222-4230
25. Coughlan MP, Mayer F (1992) The cellulose-decomposing bacteria and their enzyme
systems. In The Prokaryotes: a handbook on the biology of bacteria. 2ns edn., p. 460516. Edited by Balows A, Trper HG, Dworkin M, Harder W, Schleifer KH. SpringerVerlag, New York
26. Dees C, Ringelberg D, Scott TC, Phelps TJ (1995) Characterization of the cellulose
degrading bacterium NCIMB 10462. Appl Biochem Biotechnol 51: 263-274
27. Ding SY, Bayer EA, Steiner D, Shoham Y, Lamed R (1999) A novel cellulosomal
scaffoldin from Acetivibrio cellulolyticus that contains a family 9 glycosyl hydrolase. J
Bacteriol 181: 6720-6729
28. Elberson MA, Malekzadeh F, Yazdi MT, Kameranpour N, Noori-Dloii MR, Matte MH,
Shahamat M, Colwell RR, Sowers KR (2000) Cellulomonas persica sp. nov.
and Cellulomonas iranensis sp. nov., mesophilic cellulose-degrading bacteria isolated
from forest soil. Int J System Evol Microbiol 50: 993-996
29. Eppard M, Krumbein WE, Koch C, Rhiel E, Staley JT, Stackebrandt E (1996)
Morphological, physiological, and molecular characterization of Actinomycetes isolated
from dry soil, rocks, and monument surfaces. Arch Microbiol. 166:12-22
30. Fields MW, Mallik S, Russell JB (2000) Fibrobacter succinogenes S85 ferments ballmilled cellulose as fast as cellobiose until cellulose surface area is limiting. Appl Microbiol
Biotechnol 54: 570-574
31. Garrity GM ed. (2001) Bergeys manual of systematic bacteriology. 2 nd ed.. SpringerVerlag New York
32. Hgerdahl B, Harris H, Pye EK (1979) Association of beta-glucosidase with intact cells
ofThermoactinomyces.
Biotechnol Bioeng 21: 345-55
33. Jin F, Toda K (1989) Purification and characterization of cellulases from Clostridium
thermocopriae sp. nov. JT3-3. J Ferment Bioeng 67: 8-13

34. Kakiuchi M, Isui A, Suzuki K, Fujino T, Fujino E, Kimura T, Karita S, Sakka K, Ohmiya K
(1998) Cloning and DNA sequencing of the genes encoding Clostridium josui scaffolding
protein CipA and cellulase CelD and identification of their gene products as major
components of the cellulosome. J Bacteriol 180: 4303-4308
35. Kelly WJ, Asmundson RV, Hopcroft DH (1987) Isolation and characterization of a strictly
anaerobic, cellulolytic spore former: Clostridium chartatabidum sp. nov. Arch Microbiol
147: 169-173
36. Khan AW, Meek E, Sowden LC, Colden JR (1984) Emendation of the
genus Acetivibrio and description ofAcetivibrio cellulosolvens sp. nov., a nonmotile
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37. Kim CH (1995) Characterization and substrate specificity of an endo--1,4-D-glucanase I
(Avicelase I) from an extracellular multienzyme complex of Bacillus circulans. Appl
Environ Microbiol 61: 959-965
38. Lamed R, Naimark J, Morgenstern E, Bayer EA (1987) Specialized surface structure in
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40. Lednicka D, Mergaert J, Cnockaert MC, Swings J (2000) Isolation and identification of
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Microbiol. 23:292-299
41. Li X, Chen H, Ljungdahl L (1997) Appl Environ Microbiol 63: 4721-4728
42. MacKenzie CR, Bilous D, Johnson KG (1984) Purification and characterization of an
exoglucanase fromStreptomyces flavogriseus. Can J Microbiol 30: 1171-1178
43. Mndez BS, Pettinari MJ, Ivanier SE, Ramos CA, Sineriz F (1991) Clostridium
thermopapyrolyticum sp. nov., a cellulolytic thermophile. Int J Syst Bacteriol 41: 281-283
44. Mladenovska Z, Mathrani IM, Ahring BK (1995) Isolation and characterization
of Caldicellulosiruptor lactoaceticus sp. nov., an extremely thermophilic, cellulolytic
anaerobic bacterium. Arch Microbiol 163: 223-230
45. Mullings R, Parish JH (1984) Mesophilic aerobic Gram negative cellulose degrading
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46. Ohara H, Karita S, Kimura T, Sakka K, Ohmiya K (2000) Characterization of the
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47. Pags S, Gal L, Blaich A, Gaudin C, Tardif C, Blaich JP (1997) Role of scaffolding
protein CipC ofClostridium cellulolyticum in cellulose degradation. J Bacteriol 179: 28102816
48. Palop ML, Valles S, Pinaga F, Flors A (1989) Isolation and characterization of an
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68-71

49. Perito B, Hanhart E, Irdani T, Iqbal M, McCarthy AJ, Mastromei G (1994) Characterization
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50. Pohlschrder M, Canale-Parola E, Leschine SB (1995) Ultrastructural diversity of the
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56. Shoseyov O, Doi RH (1990) Essential 170-kDa subunit for degradation of crystalline
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thermophilum gen. nov., sp. nov.: an extremely thermophilic cellulolytic eubacterium
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59. Tamaru Y, Karita S, Ibrahim A, Chan H, Doi RH (2000) A large gene cluster for
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