The contribution of species richness and composition to bacterial services


Bacterial communities provide important services. They break down pollutants, municipal waste and ingested food, and they are the primary means by which organic matter is recycled to plants and other autotrophs. However, the processes that determine the rate at which these services are supplied are only starting to be identified. Biodiversity influences the way in which ecosystems function1, but the form of the relationship between bacterial biodiversity and functioning remains poorly understood. Here we describe a manipulative experiment that measured how biodiversity affects the functioning of communities containing up to 72 bacterial species constructed from a collection of naturally occurring culturable bacteria. The experimental design allowed us to manipulate large numbers of bacterial species selected at random from those that were culturable. We demonstrate that there is a decelerating relationship between community respiration and increasing bacterial diversity. We also show that both synergistic interactions among bacterial species and the composition of the bacterial community are important in determining the level of ecosystem functioning.

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Figure 1: The relationship between species richness and ecosystem functioning.
Figure 2: Relationship between manipulated species richness ( R ) and ecosystem functioning ( F ) over 28 days.
Figure 3: Relationship between manipulated species richness ( R ) and ecosystem functioning ( F ) over each of the three time periods.
Figure 4: Linear model coefficients as a function of the theoretical quantiles of the normal distribution.


  1. 1

    Loreau, M. et al. Biodiversity and ecosystem functioning: Current knowledge and future challenges. Science 294, 804–808 (2001)

    ADS  CAS  Article  Google Scholar 

  2. 2

    Loreau, M. & Hector, A. Partitioning selection and complementarity in biodiversity experiments. Nature 412, 72–76 (2001)

    ADS  CAS  Article  Google Scholar 

  3. 3

    Emmerson, M. C., Solan, M., Emes, C., Paterson, D. M. & Raffaelli, D. Consistent patterns and the idiosyncratic effects of biodiversity in marine ecosystems. Nature 411, 73–77 (2001)

    ADS  CAS  Article  Google Scholar 

  4. 4

    Hooper, D. U. et al. Effects of biodiversity on ecosystem functioning: A consensus of current knowledge. Ecol. Monogr. 75, 3–35 (2005)

    Article  Google Scholar 

  5. 5

    Garland, J. L. & Lehman, R. M. Dilution/extinction of community phenotypic characters to estimate relative structural diversity in mixed communities. FEMS Microbiol. Ecol. 30, 333–343 (1999)

    CAS  Article  Google Scholar 

  6. 6

    Griffiths, B. S. et al. An examination of the biodiversity–ecosystem function relationship in arable soil microbial communities. Soil Biol. Biochem. 33, 1713–1722 (2001)

    CAS  Article  Google Scholar 

  7. 7

    Franklin, R. B., Garland, J. L., Bolster, C. H. & Mills, A. L. Impact of dilution on microbial community structure and functional potential: Comparison of numerical simulations and batch culture experiments. Appl. Environ. Microbiol. 67, 702–712 (2001)

    CAS  Article  Google Scholar 

  8. 8

    Yin, B., Crowley, D., Sparovek, G., De Melo, W. J. & Borneman, J. Bacterial functional redundancy along a soil reclamation gradient. Appl. Environ. Microbiol. 66, 4361–4365 (2000)

    CAS  Article  Google Scholar 

  9. 9

    Fierer, N., Schimel, J. P. & Holden, P. A. Influence of drying-rewetting frequency on soil bacterial community structure. Microb. Ecol. 45, 63–71 (2003)

    CAS  Article  Google Scholar 

  10. 10

    Cavigelli, M. A. & Robertson, G. P. The functional significance of denitrifier community composition in a terrestrial ecosystem. Ecology 81, 1402–1414 (2000)

    Article  Google Scholar 

  11. 11

    Cavigelli, M. A. & Robertson, G. P. Role of denitrifier diversity in rates of nitrous oxide consumption in a terrestrial ecosystem. Soil. Biol. Biochem. 33, 297–310 (2001)

    CAS  Article  Google Scholar 

  12. 12

    Naeem, S., Hahn, D. R. & Schuurman, G. Producer–decomposer co-dependency influences biodiversity effects. Nature 403, 762–764 (2000)

    ADS  CAS  Article  Google Scholar 

  13. 13

    Griffiths, B. S. et al. The relationship between microbial community structure and functional stability, tested experimentally in an upland pasture soil. Microb. Ecol. 47, 104–113 (2004)

    CAS  Article  Google Scholar 

  14. 14

    Horz, H. P., Barbrook, A., Field, C. B. & Bohannan, B. J. M. Ammonia-oxidizing bacteria respond to multifactorial global change. Proc. Natl. Acad. Sci. USA 101, 15136–15141 (2004)

    ADS  CAS  Article  Google Scholar 

  15. 15

    Huston, M. A. Hidden treatments in ecological experiments: Re-evaluating the ecosystem function of biodiversity. Oecologia 110, 449–460 (1997)

    ADS  Article  Google Scholar 

  16. 16

    Kitching, R. L. An ecological study of water-filled tree-holes and their position in the woodland ecosystem. J. Anim. Ecol. 40, 281–302 (1971)

    Article  Google Scholar 

  17. 17

    Bell, T. et al. Larger islands house more bacterial taxa. Science 308, 1884 (2005)

    CAS  Article  Google Scholar 

  18. 18

    van der Heijden, M. G. A. et al. Mycorrhizal fungal diversity determines plant biodiversity, ecosystem variability and productivity. Nature 396, 69–72 (1998)

    ADS  CAS  Article  Google Scholar 

  19. 19

    McGrady-Steed, J., Harris, P. M. & Morin, P. J. Biodiversity regulates ecosystem predictability. Nature 390, 162–165 (1997)

    ADS  CAS  Article  Google Scholar 

  20. 20

    Tilman, D. et al. Diversity and productivity in a long-term grassland experiment. Science 294, 843–845 (2001)

    ADS  CAS  Article  Google Scholar 

  21. 21

    Hector, A. et al. Plant diversity and productivity experiments in European grasslands. Science 286, 1123–1127 (1999)

    CAS  Article  Google Scholar 

  22. 22

    Curtis, T. F., Head, I. M. & Graham, D. Theoretical ecology for engineering biology. Environ. Sci. Technol. 37, 64A–70A (2003)

    ADS  Article  Google Scholar 

  23. 23

    Cardinale, B. J., Palmer, M. A. & Collins, S. L. Species diversity enhances ecosystem functioning through interspecific facilitation. Nature 415, 426–429 (2002)

    ADS  CAS  Article  Google Scholar 

  24. 24

    Zengler, K. et al. Cultivating the uncultured. Proc. Natl. Acad. Sci. USA 99, 15681–15686 (2002)

    ADS  CAS  Article  Google Scholar 

  25. 25

    Rappe, M. S. & Giavannoni, S. J. The uncultured microbial majority. Annu. Rev. Microbiol. 57, 369–394 (2003)

    CAS  Article  Google Scholar 

  26. 26

    Schimel, J. P. & Gulledge, J. Microbial community structure and global trace gases. Global Change Biol. 4, 745–758 (1998)

    ADS  Article  Google Scholar 

  27. 27

    Sasser, M. MIDI Inc. technical note 101: Identification of bacteria by gas chromatography of cellular fatty acids (Newark, 2001).

  28. 28

    Thompson, I. P., Bailey, M. J., Ellis, R. J. & Purdy, K. J. Subgrouping of bacterial-populations by cellular fatty-acid composition. FEMS Microbiol. Ecol. 102, 75–84 (1993)

    CAS  Article  Google Scholar 

  29. 29

    Cohan, F. M. What are bacterial species? Annu. Rev. Microbiol. 56, 457–487 (2002)

    CAS  Article  Google Scholar 

  30. 30

    Zibilske, L. M. Methods of Soil Analysis, Part 2. Microbiological and Biochemical Properties 835–863 (SSSA, Madison, 1994)

    Google Scholar 

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We are grateful to J. Fryxell, T. Nudds and their graduate students for providing comments on the original manuscript, to A. Singer for help developing the technique to measure bacterial respiration, and the Centre for Ecology and Hydrology in Oxford for providing the laboratory space. T. B. was supported by Fonds quebecois de la recherche sur la nature et les technologies, the Natural Sciences and Engineering Council of Canada, and the Clarendon Fund (Oxford University).Author contributions The experiment was originally conceived by T.B., J.A.N. and A.K.L. The laboratory work was conducted by T.B. with the help of A.K.L. and S.L.T. The experimental design was conceived by A.K.L. and developed by T.B., J.A.N. and B.W.S. The statistical analyses were performed by B.W.S. and T.B. The manuscript was written principally by T.B. with extensive input from J.A.N., B.W.S. and A.K.L.

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Correspondence to Andrew K. Lilley.

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Supplementary information

Supplementary Discussion

This file contains discussion of additional analysis of the experiment in which we take a detailed look at the interaction between species composition and species richness. (PDF 19 kb)

Supplementary Table S1

This files contains a list of the species level identities used in the experiment. (PDF 13 kb)

Supplementary Table S2

This files contains an outline of the experiment design. (PDF 15 kb)

Supplementary Figure S1

This file contains a figure showing the relationship between the effect of each species on respiration and sensitivity of that effect to the level of diversity. (PDF 19 kb)

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Bell, T., Newman, J., Silverman, B. et al. The contribution of species richness and composition to bacterial services. Nature 436, 1157–1160 (2005).

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