MICROBIAL

BIODIVERSITY
Devon J. Bradleynw and Jennifer B. H. Martinyw
n
Brown University and University of California

I. Introduction polymerase chain reaction (PCR) A method for

II. Characterizing Microbial Diversity copying DNA sequences by repeated cycles of syn-
III. What is Microbial Diversity? thesis using specific primers and DNA polymerase.
IV. Maintenance of Microbial Diversity virus A genetic element containing either DNA or
V. Importance for Ecosystems RNA that replicates in cells but is characterized by
having an extracellular state.

GLOSSARY
THE OVERWHELMING MAJORITY of biological
diversity is microbial. Microorganisms span all three
archaea One of the two entirely microbial domains
domains of life and are typically defined as unicellular
of single-celled organisms, evolutionarily distinct
life forms that can only be observed with a micro-
from bacteria. Many archaeal characteristics are
scope, including bacteria, archaea, viruses, and many
more similar to eukaryotes than bacteria.
unicellular eukaryotes (e.g., some fungi and protists).
bacteria An entirely microbial domain of single-celled
This article asks three fundamental questions about
organisms, evolutionarily distinct from Archaea.
microbial diversity: how do we measure it? how much
eukarya A primarily microbial domain of organisms
is there? what is its role in ecosystems? To address
having a membrane-enclosed nucleus and other
these questions we discuss the methodologies em-
organelles; includes animals, plants, fungi, and
ployed to characterize microbial diversity, we survey
protists.
the magnitude of global microbial diversity, and review
fungi Nonphototrophic, heterotrophic eukaryotic
some of the ecological and evolutionary forces thought
microorganisms that contain rigid cell walls; in-
to be important in maintaining microbial diversity. Fi-
cludes mushrooms, molds, and the fungal part of
nally, we discuss what is known about the role of mi-
lichens.
crobial diversity in ecosystem functioning and
metagenomics The genomic analysis of microorgan-
consider how microbes mediate ecosystem function-
isms by direct extraction and cloning of community
ing in the face of human-driven global changes.
DNA from an environmental sample.
microorganism Single-celled organisms that can
only be observed with a microscope, including
bacteria, archaea, small eukaryotes, and viruses. I. INTRODUCTION
operational taxonomic unit (OTU) A group of
organisms regarded as being distinct from other Although it might not be immediately obvious, our
groups, based on any clearly defined variables. world is a microbial one. Biodiversity is usually
Encyclopedia of Biodiversity
Copyright & 2007 Elsevier Inc. All rights of reproduction in any form reserved. 1
2 _______________________________________________________________________________________________________________________________________________________________________________________________________________________________________________ M I C R O B I A L B I O D I V E R S I T Y _______________________________________________________________________________________________________________________________________________________________________________________________________________________________________________

discussed in terms of large organisms, but no organ- Prior to the development of molecular methods, it
isms are more ubiquitous, abundant, or diverse than was particularly challenging to understand the
microorganisms. Microorganisms were the first cellu- evolutionary relationships between organisms. It was
lar life forms and were active more than 3 billion years generally accepted that there were two basic kinds of
prior to the appearance of macroorganisms. The met- organisms, prokaryotes and eukaryotes, distinguished
abolic activities that they carried out during this time by the absence or presence of a membrane-bound nu-
were critical for creating the conditions for the evolut- cleus. In the mid-1970s, Woese and colleagues assem-
ion of multicellular forms. The universal tree of life bled the universal tree of life based on sequence
(Fig. 1) emphasizes this point; multicellular Eukarya information of small-subunit ribosomal RNA (SSU
are crown groups compared with the deeply rooted rRNA). This breakthrough revealed that life consisted
Bacteria and Archaea. Microbes force us to stretch our of two very distinct microbial domains, the Bacteria
imagination about the limits of metabolic lifestyles, the and Archaea, which appear to have diverged very early
geography of life, and the roles that organisms play in on in evolutionary history, and a third mostly micro-
our lives. bial domain, Eukarya, the exact origins of which
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FIGURE 1 The universal tree of life showing the position of Archaea and Bacteria
relative to Eucarya. The placement of organisms on this phylogenetic tree is based on
the analyses of SSU rRNA sequeunces of organisms from within each domain. From
Pace (1999), with permission.
_____________________________________________________________________________________________________________________________________________________________________________ M I C R O B I A L B I O D I V E R S I T Y __________________________________________________________________________________________________________________________________________________________________________ 3

remain obscure. Archaea and Bacteria share similar many levels of biological organization from the alleles
morphological characteristics but Archaea possess at a particular genetic locus to community composi-
several cellular and genetic characteristics that are tion among biomes.
more similar to eukaryotes and these organisms tend
to be more abundant under extreme environments. As
a consequence, Bacteria and Archaea often perform A. Culture Techniques
quite different roles. For example, photosynthesis is
Throughout most of the twentieth century, micro-
performed by Bacteria, spread across five different
biologists used culturing techniques to characterize
clades, and is not found among Archaea, while
the composition of microbial communities. Tradition-
methanogenesis is restricted to Archaea. Members of
ally, these techniques involve trying to grow micro-
these two domains occupy all habitats on Earth.
organisms from environmental samples on artificial
media and mimicking natural environmental condi-
tions, such as temperature and light, in a laboratory
II. CHARACTERIZING MICROBIAL setting. These traditional cultivation techniques tend
to select for fast-growing taxa, and therefore grossly
DIVERSITY underestimate actual diversity, but more importantly,
traditional cultivation techniques are thought to cap-
Although delineating species can be challenging for
ture less than 1% of all bacterial species (Ward et al.,
some macroorganisms, defining a microbial species is
1990), and therefore provide a biased view of micro-
particularly challenging, in part because of their gene-
bial community composition in most environments. In
tics. For sexual organisms, a species is usually defined
the early twentieth century, however, promising new
as a group of potentially interbreeding organisms. But
cultivation techniques were developed to overcome
many microorganisms, bacteria in particular, are asex-
these deficiencies. For instance, Kaeberlein et al.
ual. In addition, microorganisms are too small to de-
(2002) used small diffusion chambers to grow bacte-
fine by morphological characteristics, as is usually
rial colonies in a more natural environment and re-
done by plant and animal taxonomists. Thus, in stark
source regimen. Cho and Giovannoni (2004) used
contrast to diversity surveys of large organisms, these
high-throughput methods to culture novel strains of
microorganisms often focus on variation in their DNA
oceanic bacteria in low-nutrient media. These newer
rather than in their phenotype. For instance, bacterial
techniques have allowed microbial biologists to study
species are defined as a group of strains that share
the ecophysiological properties of organisms that had
more than 70% of their genome. Operationally, this
been previously inaccessible.
definition is applied by melting the genomic DNA of
two bacteria and measuring the rate at which they re-
anneal, a rate determined in part by the similarity of
the organisms’ genomes.
B. Genetic Methodologies
For microbial community studies, the 70% genomic The development of culture-independent methods,
similarity definition is impractical to apply. Instead, based on the genetic characterization of microbial
researchers use genetic markers to characterize micro- communities, has allowed the detection of noncultur-
bial community composition and define an operational able species and a more detailed description of micro-
taxonomic unit (OTU). For instance, in studies that bial communities. Because rDNA molecules are found
sequence the 16S rDNA of bacteria (see below), a in all organisms and are probably rarely laterally trans-
common OTU definition is a group of sequences that ferred, they have become a standard phylogenetic
are similar by 97%. This definition is based on the marker of microorganisms. Pace et al. (1986) built
rough relationship between 16S rDNA similarity and upon this idea and used the polymerase chain reaction
the 70% genomic similarity definition (Stackebrandt (PCR) to amplify rDNA genes from community DNA
and Goebel, 1994). Once microbial species, or other in environmental samples. After PCR, the products can
taxonomic units, are defined, then the diversity of a then be further assayed to characterize the diversity of
microbial community can be measured. In its simplest the community rDNA genes. Current open-access data-
form, community diversity is the richness (total bases such as GenBank and the Ribosomal Database
number of OTUs) and evenness (relative abundance Project contain thousands of rDNA gene sequences.
of the OTUs) in a defined area. A full description of Numerous methods can be used to characterize
microbial diversity, however, includes variation at the diversity of PCR-amplified fragments. Cloning
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followed by DNA sequencing provides the most de- TABLE I

tailed information, but is also expensive. ‘‘Fingerprinting’’ Comparison of the total number of species currently described and
methods are a cheaper alternative for characterizing the number of ‘‘species’’ predicted to exist. The term ‘‘species’’ is
microbial community composition. For instance, used loosely as it must be defined differently across taxa
terminal restriction fragment length polymorphism Group Number of ‘‘species’’
(T-RFLP), a modification of the conventional RFLP
Described Estimated % Described
approach, allows the characterization of a community
6 6
by analyzing the size polymorphisms of PCR-amplified Eukaryotes 1.8  10 12  10 15.0
genes that have been cut at precise sequence locations Fungi 7.2–10  104 1.5  106 5.3
Archaea 217
by restriction enzymes. The resulting fragments are
Bacteria 5007 4.5–1000  106 0.1
separated by gel electrophoresis and produce a ban- Viruses 1550 4.0  105 0.4
ding pattern that can be used as a fingerprint for com-
paring the similarity between microbial community Modified from Martiny and Field (2005). The fungal estimates are
composition among samples. from Hawksworth et al. (1995) and Rossman (1995).
As DNA sequencing has become faster and cheaper,
the field of metagenomics has emerged. Metagenomics have tried to estimate microbial diversity globally or
uses the DNA sequences of fragments of DNA obtained within particular habitat types.
from a natural microbial community to describe the
collective diversity of microbial genomes. Shotgun se-
quencing breaks up an organism’s or a community’s
DNA into a myriad of short fragments that are
individually cloned and sequenced. These fragments
A. Bacterial and Archaeal Diversity
are then computationally ordered based on overlaps in The sheer abundance of bacteria makes estimating the
the genetic code and reassembled into larger genomic number of bacterial taxa difficult. Curtis and Sloan
fragments. For instance, Venter et al. (2004) applied (2005) calculate that there are more bacteria on Earth
this approach to an open ocean environment in the than there are stars in the universe and that most
Sargasso Sea. This pilot project detected 150 new spe- microbial communities harbor 1010–1017 bacterial
cies of bacteria and over 1.2 million new genes. cells that compose greater than 107 different taxo-
Screening of metagenomic soil libraries has already led nomic groups and innumerable functional groups
to the identification of various novel biomolecules in- (Falkowski, 2002). The magnitude of bacterial diver-
cluding enzymes and antibiotics. An alternative ap- sity is still a matter of debate and perhaps even beyond
proach allows the insertion of large continuous practical calculation. Dykhuizen (1998) has specu-
sequences of DNA to be incorporated into bacteria to lated that there could be 10 billion ‘‘species’’ of bacteria
form artificial chromosomes (BACs), which are then on Earth. It has only recently been demonstrated that
sequenced. This latter approach is much more labor- different habitats harbor different amounts of pro-
intensive and time consuming, but allows one to see karyotic diversity (Fig. 2). For instance, Torsvik et al.
how genes are organized within microbes, which is (2002) used DNA–DNA reassociation methods to
important information if one is interested in under- quantify the total genomic diversity in different com-
standing their evolutionary history. munities. They estimate that a few grams of soil or
sediment contain thousands of bacterial and archaeal
‘‘species.’’ In comparison, aquatic environments appear
III. WHAT IS MICROBIAL DIVERSITY? to support fewer bacterial taxa (Torsvik et al., 2002;
Curtis and Sloan, 2005).
The immense diversity, small size, and clonal nature of Because of their high taxonomic and evolutionary
most microorganisms explain why quantifying the diversity, bacteria are incredibly functionally diverse.
biodiversity of microorganisms is fundamentally dif- Bacteria have a striking variety of modes of energy
ferent from quantifying that of macroorganisms. conversion, many unique metabolic pathways, and can
Microorganisms are so abundant and diverse that only utilize a wide range of substrates. Bacteria can oxidize
a minute fraction of their diversity has been described and reduce inorganic sulfur and can assimilate atmos-
(Table I). Currently, it is infeasible to completely pheric nitrogen. Many bacteria have an anaerobic
quantify the microbial diversity in even a gram of soil energy metabolism and are often the only organisms
or a liter of seawater. However, a number of studies living in anoxic environments. Bacteria can even
_____________________________________________________________________________________________________________________________________________________________________________ M I C R O B I A L B I O D I V E R S I T Y __________________________________________________________________________________________________________________________________________________________________________ 5

4.5

Genomic equivalents (log scale)

3.5

2.5

1.5

0.5

0
Marine Pasture Forest Arable Marine Forest Salt pond
pristine soil soil soil fish-farm cultivated
sediment sediment bacteria

FIGURE 2 Prokaryotic diversity (genome equivalents) estimated from the reassociation rate of
DNA isolated from various habitats. Estimates for pasture soil and arable soil are averages.
Modified from Torsvik et al. (2002).

degrade xenobiotic compounds, such as herbicides archaea are by far the more abundant ammonia oxi-
and pesticides, which are toxic to other organisms. dizers in both pristine and agricultural soils.
Because archaea require particular effort to culture,
the diversity of this group is historically less well
understood than bacteria. Molecular phylogenetic
B. Microbial Eukaryotic Diversity
surveys, more commonly applied to the study of bac- Molecular methodologies that have revolutionized the
terial communities, are now more frequently being understanding of bacterial and archaeal diversity have
used to characterize the genetic diversity and ecolo- only recently been adapted to study eukaryotic diver-
gical significance of archaea in a variety of environm- sity. Perhaps not surprisingly, recent molecular surveys
ents including hot springs, deep-sea hydrothermal suggest that morphological characteristics may con-
vents, and coastal waters. Approximately 217 archaeal ceal a large amount of eukaryotic diversity. For exam-
‘‘species’’ have been described, compared with over ple, Slapeta et al. (2005) analyzed the rDNA genes of
5000 bacterial ‘‘species,’’ and we lack estimates of how protists in two ponds and found high within-morpho-
diverse these prokaryotes might be. type diversity. In addition, they detected a number of
Archaea have typically been found living at physical sequences very divergent from known protist species,
or chemical extremes such as high temperature, high implying that the description of global protist diversity
salinity, or strictly anaerobic environments and are is far from complete. Finlay and Fenchel (2004) argue
immensely physiologically and metabolically diverse. that while local microbial eukaryotic diversity is high,
Methanogenic archaea, which occupy anaerobic hab- global diversity may be relatively low. They document
itats such as sediments, marshes, and hydrothermal that many free-living protists (as defined by morpho-
vents, produce methane as a by-product of their me- logy) are cosmopolitan and suggest that nearly all free-
tabolism, a process restricted to this domain. Recently, living ciliates have already been discovered.
archaea are being found in more benign marine, fresh- Like protists, fungal species have been traditionally
water, and soil environments. For example, it has defined morphologically. Hawksworth (1991) suggests
generally been assumed that specialist bacteria, which that there are at least 1.5 million fungal species glob-
convert ammonia into nitrate, a key step in nitrifica- ally. Somewhere between 72,000 and 100,000 fungal
tion, are the main ammonia oxidizers in soil. Surpri- species have been described, suggesting that as little as
singly, Leininger et al. (2006) recently revealed that 5% of fungal diversity has been discovered. Moreover,
6 _______________________________________________________________________________________________________________________________________________________________________________________________________________________________________________ M I C R O B I A L B I O D I V E R S I T Y _______________________________________________________________________________________________________________________________________________________________________________________________________________________________________________

with the application of genetic methods such as those exceeds the rate of extinction. If laboratory studies are
applied to bacteria and archaea, estimates of global indicative (52 or 53), the potential for rapid diversifi-
fungal diversity will likely increase substantially. For cation seems greater in microorganisms than macro-
example, Vandenkoornhuyse et al. (2002) surveyed organisms. Bacteria and archaea, in particular, have
the genetic diversity of fungi in the roots of one grass several traits that make rapid speciation likely. Because
species and found 49 different fungal OTUs, only they exist in large populations and have rapid growth
seven of which were similar to known DNA sequences, rates, they have the capacity to accumulate large num-
and only 6% of which have known ecological roles. bers of novel mutations, which can generate substan-
tial genetic variation. They can also readily take up
DNA and get rid of extraneous DNA. Gene transfer
C. Viral Diversity and recombination may also accelerate diversification
Given the remarkable diversity of bacteria, archaea, further. Less is known about the extinction rates of
and microbial eukaryotes, it is perhaps not surprising microbes relative to larger organisms; however, many
that the viruses that infect them appear to be similarly researchers have posited that extinction rates of mi-
abundant and diverse. Viral abundance is estimated to crobial taxa may be low because many microorganisms
be on the order of 1031, the majority of which infect have large population sizes and have an ability to
bacteria and archaea. One kilogram of marine sediment survive in a range of environments and tolerate harsh
may harbor a million different viral genotypes, and conditions.
200 l of seawater contain more than 5000 viral geno- From an ecological perspective, once this diversity
types. In general, viral abundance is well correlated is created, the question remains how this incredible
with bacterial abundance, and in fact, bacteriophages evolutionary diversity coexists in contemporary com-
are estimated to infect and kill between 4% and 50% of munities. Horner-Devine et al. (2003a) review eco-
bacteria produced daily in aquatic environments. How, logical factors, known to be of importance to plant and
or even whether, viral diversity varies by habitat type is animal communities, which may also influence the
unclear. Breitbart and Rohwer (2005) reviewed the diversity of microorganisms. Below we discuss four of
extent and distribution of viral diversity in marine these factors, and review recent studies that test
habitats and proposed that viruses are highly diverse whether these factors are important for environmen-
on a local scale but that this diversity does not vary tal microbial communities.
much over space. In other words, local viral diversity
may be representative of global viral diversity.
A. Environmental Heterogeneity
Among large organisms, species diversity increases
IV. MAINTENANCE OF MICROBIAL with the heterogeneity of habitat types available. For
DIVERSITY instance, bird species diversity increases with foliage
height diversity and mite diversity increases with the
Ecologists have long been motivated to understand the number of habitat types in soil subhorizons. Both lab-
factors that influence the extraordinary diversity of life oratory and field studies support the hypothesis that
on Earth. However, despite the extent of microbial increased environmental heterogeneity increases bac-
diversity, most of what we know about the mainte- terial diversity. For example, spatial isolation in the
nance of diversity is based on studies of plants and soil matrix, created by low moisture content, plays an
animals. For example, Hutchinson (1961) observed important role in determining community composi-
two species of water-bugs in a homogenous aquatic tion. Zhou et al. (2002) observed that soils saturated
environment and wondered how they could coexist if with water had fewer bacterial taxa and a less even
they both use similar nutrient resources; the same is distribution of taxa than patchy, unsaturated soils.
true about the thousands of microbial taxa that coexist Treves et al. (2003) examined the competitive dynam-
in all habitats. Although a synthetic view of microbial ics of two species growing on a single resource in a
diversity might not be possible, it is useful to consider uniform sand matrix under varied moisture content.
whether universal patterns and processes govern the One species dominated the community under satu-
diversity of all organisms, large and small. rated conditions, which were also highly connected,
From an evolutionary perspective, high microbial suggesting that these conditions allow competitive in-
diversity is created because the rate of diversification teractions to structure the community. As moisture
_____________________________________________________________________________________________________________________________________________________________________________ M I C R O B I A L B I O D I V E R S I T Y __________________________________________________________________________________________________________________________________________________________________________ 7

content and connectivity decreased, the less-compet- fluorescens over a wide range of nutrient concentra-
itive species became established in the community. tions and found that diversity peaked at intermediate
productivities. However, Benlloch et al. (1995) sur-
veyed two lagoons with different levels of primary
B. Disturbance productivity and found a greater number of unique
bacterial rRNA gene sequences in the more productive
Most communities are subject to different types of
lagoon, suggesting that bacterial taxonomic richness
disturbances that vary in both frequency and intensity.
increases with productivity. Finally, Horner-Devine
Events that disrupt the physical environment or alter
et al. (2003b) estimated bacterial taxonomic richness
resource and substrate availability are known to influ-
along a primary productivity gradient in freshwater
ence plant and animal community diversity. Across a
mesocosms. Although overall bacterial richness was
wide range of taxa, including corals, butterflies, plank-
not significantly related to productivity, productivity
ton, and tropical trees, diversity peaks at intermediate
did influence the richness of separate taxonomic
intensities or frequencies of small-scale disturbances.
groups. For example, some bacterial groups exhibited
For example, algal diversity on intertidal boulders is
a significant hump-shaped relationship with pro-
highest when waves overturn the boulders, exposing
ductivity while other groups, such as the a-Proteobac-
new substrate for colonization.
teria, exhibited a U-shaped relationship with primary
It is difficult to test whether the intermediate dis-
productivity.
turbance hypothesis explains patterns of microbial
diversity in the wild because disturbance is typically
confounded with other factors such as plant cover or D. Dispersal
soil structure. Field and laboratory studies that have
examined the relationship between disturbance and The small size and high abundance of microorganisms
bacterial diversity have generally found that the inter- have led microbiologists to generalize that microbial
mediate disturbance hypothesis holds. For example, dispersal rates to new habitats is very high, overwhel-
Bruce et al. (1995) examined the diversity of mercury- ming spatial differences in taxon abundance. Recently,
resistance genes in sediment-associated bacteria at this view has been called into question, as many stud-
sites with varying levels of mercury contamination. ies demonstrate biogeographic differences among mi-
Consistent with predictions of the intermediate dis- crobial communities (Martiny et al., 2006). This
turbance hypothesis, sites exposed to intermediate evidence suggests that microbial diversity might also
levels of mercury had the highest genetic diversity and be influenced by dispersal and isolation patterns
both pristine and heavily contaminated sites had low (Hubbell, 2001). Based on this theory, Sloan et al.
diversity. Walsh et al. (2005) compared archaeal 16S (2006) argue that local bacterial community compo-
rRNA gene diversity along a salinity gradient prone to sition is in part a product of stochastic dispersal
high-salinity fluctuations. Heavy-rainfall events im- events, rather than solely niche differentiation among
pose local high mortality of haloarchaea, and this dis- taxa (as is suggested by the three factors above).
turbance regime appears to promote archaeal diversity. The importance of dispersal relative to other factors
influencing microbial diversity requires further
investigation.
C. Productivity
Productivity, the rate at which energy flows through an
ecosystem, has long been considered a primary deter- V. IMPORTANCE FOR ECOSYSTEMS
minant of both plant and animal diversity. A current
ecological paradigm is that peak diversity occurs at Microorganisms are the drivers of ecosystem proc-
intermediate levels of productivity, even though this esses; particularly, decomposition, nutrient minerali-
relationship appears to be scale-dependent. zation, and trace gas emission and consumption.
A number of studies consider the relationship be- Microorganisms are intimately involved in critical
tween primary productivity and microbial diversity. pathways of biogeochemical cycles and are the major
Productivity appears to be related to microbial diver- pathways by which some elemental forms are regene-
sity; in some instances, these patterns resemble those rated to be used by other organisms.
of plants and animals. For example, Kassen et al. For instance, microbes have a central role in almost
(2000) grew cultures of the bacterium Pseudomonas all aspects of the nitrogen cycle. Nitrogen fixation is
8 _______________________________________________________________________________________________________________________________________________________________________________________________________________________________________________ M I C R O B I A L B I O D I V E R S I T Y _______________________________________________________________________________________________________________________________________________________________________________________________________________________________________________

the process by which atmospheric nitrogen is conver- both strongly interacted with temperature and precip-
ted into ammonia, which is quickly ionized to ammo- itation. Methane-oxidizing bacteria showed a unique
nium, a form of nitrogen usable by plants. This response to increased precipitation and temperature;
process, carried out by nitrogen-fixing bacteria, is es- the relative abundance of common methanotroph cla-
sential for maintaining soil fertility and agricultural des increased in response to both variables, while
productivity. novel clades had varied responses. Two of the novel
If an entire functional group, such as the nitrogen- clades did not respond to these environmental
fixing bacteria, were removed from an ecosystem, the changes, while one novel clade increased in relative
resulting changes would dramatically alter resource abundance to increased temperature and precipitation.
supply, processing rates, and plant productivity. How- Finally, several studies illustrate the potential impor-
ever, below this broad functional group classification, tance of feedbacks between microbial and plant com-
it remains unclear whether microbial diversity matters munities that could alter the overall ecosystem response
for ecosystem processes. Recently, numerous studies to global change and the potential unpredictable nature
have highlighted the incredible ‘‘microdiversity’’ of of these feedbacks. In a one-year factorial field exper-
bacteria in some habitats. For example, Acinas et al. iment, Mitchell et al. (2003) tested the influence of plant
(2004) sampled 16S rDNA sequences from a coastal diversity, elevated CO2 levels, and nitrogen deposition
bacterioplankton community. They found 516 unique on the severity of disease caused by fungal pathogens in
sequences, 50% of which fell into clusters containing a diverse grassland community. All three components of
less than 1% sequence divergence. Thus, one current global change substantially increased disease severity in
avenue of research is to understand whether this type the plant community. Additionally, fungi might serve as
of fine-scale diversity or other levels of diversity have a negative feedback under increased atmospheric CO2
consequences for ecosystem processes. because mycorrhizae sequester increased amounts of
carbon in living, dead, and residual hyphal biomass. In
contrast, nitrogen deposition could increase turnover
rates of fungal tissue and negate CO2 effects on fungal
A. Microorganisms and Global Change biomass (Treseder and Allen, 2000).
The composition of microbial communities is known A more complete understanding of the ecosystem
to vary with land-use type, temperature, agricultural impacts of global change will require understanding
growing practices, nutrient status, pollutants, and the response of the diversity of both microorganisms
other environmental variables associated with global and macroorganisms to multiple environmental vari-
change. Additionally, there is some evidence that cer- ables associated with global change.
tain properties of entire microbial communities can be
altered by global changes, such as total microbial
biomass, rates of respiration, and biogeochemical See Also the Following Articles
transformations. Little is known, however, how an- ARCHAEA, ORIGIN OF  BACTERIAL BIODIVERSITY 
thropogenic impacts will affect microbial communities, BIOGEOCHEMICAL CYCLES  MICROBIAL BIODIVERSITY,
and ultimately ecosystem functioning. MEASUREMENT OF  MICROBIAL DIVERSITY 
Several recent studies have examined the links be- MICROORGANISMS (MICROBES), ROLE OF
tween microbial composition and ecosystem function-
ing and support the hypothesis that global changes
will have important consequences for microbially me- Bibliography
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