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Microbiology is the study of microorganisms—biological entities too small to be seen with the unaided eye. Most major advances in microbiology have occurred within the past 150 years, and several important subdisciplines of microbiology have developed during this time, including microbial ecology, molecular biology, immunology, industrial microbiology, and biotechnology. Microorganisms of various types exist in all three domains of life (the Bacteria, Archaea, and Eukarya), and they are by far the most abundant life forms on Earth. Microscopic biological agents include bacteria, archaea, protists (protozoa and algae), fungi, parasitic worms (helminths), and viruses. Although a small percentage of microorganisms are harmful to certain plants and animals and may cause serious disease in humans, the vast majority of microorganisms provide beneficial services, such as assisting in water purification and the production of certain foods, and many are essential for the proper functioning of Earth’s ecosystems.
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W Matthew Sattley, Division of Natural Sciences, Indiana Wesleyan University,
Marion, Indiana, USA
Michael T Madigan, Department of Microbiology, Southern Illinois University,
Carbondale, Illinois, USA
Introductory article
Article Contents
Microbiology and its Historical Roots
Classification and Basic Characteristics of
Microbial Ecology
Medical Microbiology
Applications in Microbiology
Online posting date: 14th August 2015
Microbiology is the study of microorgan-
isms – biological entities too small to be seen
with the unaided eye. Most major advances in
microbiology have occurred within the past 150
years, and several important subdisciplines of
microbiology have developed during this time,
including microbial ecology, molecular biology,
immunology, industrial microbiology and biotech-
nology. Microorganisms of various types exist in
all three domains of life (the Bacteria,Archaea and
Eukarya), and they are by far the most abundant
life forms on Earth. Microscopic biological agents
include bacteria, archaea, protists (protozoa and
algae), fungi, parasitic worms (helminths) and
viruses. Although a small percentage of microor-
ganisms are harmful to certain plants and animals
and may cause serious disease in humans, the vast
majority of microorganisms provide beneficial
services, such as assisting in water purification
and the production of certain foods, and many
are essential for the proper functioning of Earth’s
Microbiology and its Historical
Microbiology is the study of microorganisms, microscopic organ-
isms that include in particular the bacteria, a large group of
very small cells that have enormous basic and practical signi-
cance (Madigan et al., 2015). Microbiology considers all aspects
of microbial cells, including their structure, metabolism, diver-
sity, genetics and evolution, ecology and roles in infectious dis-
eases. Microbiology is composed of several subdisciplines, each
of which is focused on part of the broader science. Scientists
eLS subject area: Microbiology
How to cite:
Sattley, W Matthew and Madigan, Michael T (August 2015)
Microbiology. In: eLS. John Wiley & Sons, Ltd: Chichester.
DOI: 10.1002/9780470015902.a0000459.pub2
who study microorganisms, called microbiologists, typically spe-
cialise in one or more of these areas (Table 1).
The science of microbiology developed later than other bio-
logical sciences, primarily because needed tools, such as the
microscope, had to be developed to convincingly prove that
microorganisms (also called microbes) exist. Following early
microscopic discoveries, methods for the culture and identi-
cation of microorganisms were developed, and from these, our
understanding of the enormous benecial and detrimental effects
of microbes began to unfold. We review some historical high-
lights in microbiology now.
The discovery of microorganisms
The English naturalist Robert Hooke (1635–1703) was an early
microscopist and published the rst book devoted entirely to
microscopic observations of microorganisms. Hooke prepared
detailed and quite accurate drawings of moulds (fungi) and many
other microbes, and these were the rst known description of
The rst person to see bacteria, which are typically much
smaller than moulds, was the Dutch amateur microscopist Antoni
van Leeuwenhoek (1632–1723). van Leeuwenhoek constructed
simple microscopes that contained a single lens and used them
to examine various natural substances. These microscopes were
crude by today’s standards, but by careful manipulation and
focusing, van Leeuwenhoek was able to see a wide variety of
microorganisms, including bacteria. van Leeuwenhoek reported
his discoveries in a series of letters to the Royal Society of Lon-
don, which were then published in the Philosophical Transac-
tions of the Royal Society, one of the most prestigious scientic
journals of the era and the rst in the world exclusively devoted to
science. His communications revealed a previously hidden micro-
bial world that existed in water, nutrient solutions, the oral cavity
and virtually anywhere one could imagine. van Leeuwenhoek’s
discoveries also boosted the long held belief that invisible agents
of some sort were the cause of infectious diseases, a belief that
was not scientically conrmed until nearly 200 years later. See
also:Leeuwenhoek, Antoni van;Light Microscopy;History
of Bacteriology
The golden age of microbiology
Major advances in microbiology in the nineteenth and early twen-
tieth centuries surrounded four major scientic questions of that
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Table 1 Selected major subdisciplines of microbiology
Subdiscipline Focus
Agricultural/soil microbiology Microbial diversity and processes in soils
Aquatic microbiology Microbial processes in water and wastewaters
Biotechnology Production of high-value products by genetically engineered microorganisms
Genomics Genome sequencing and analyses
Immunology The immune response
Industrial microbiology Large-scale production of antibiotics and commodity chemicals
Medical microbiology Nature and control of infectious diseases
Microbial biochemistry Enzymes, chemical reactions in cells, structural biology
Microbial ecology Microbial diversity and activity in natural habitats, biogeochemistry
Microbial genetics Genes, heredity and genetic variation
Microbial physiology Nutrition, metabolism and bioenergetics
Microbial systematics Classication and nomenclature
Molecular biology Nucleic acids and proteins, genetic information processing
Virology Viruses and subviral particles
period: (1) can life emerge from nonlife, (2) do microorganisms
cause infectious diseases, (3) how diverse is the microbial world
and (4) do soil and water microbes carry out any benecial activ-
ities? These questions were addressed, respectively, through the
research of four giants in the then growing eld of microbiol-
ogy: the French chemist Louis Pasteur (1822–1895), the German
physician Robert Koch (1843–1910), the Dutch microbiologist
Martinus Beijerinck (1851–1931) and the Russian microbiologist
Sergei Winogradsky (1856–1953).
Pasteur initiated studies on the mechanism of the alcoholic fer-
mentation, which in the mid-nineteenth century was assumed to
be a strictly chemical process. Through microscopic observations
and other rigorous experiments, Pasteur showed that the fermen-
tation was actually caused by the metabolic activities of yeast
cells. Pasteur then used these insights to design a series of clas-
sic experiments to disprove the theory of spontaneous generation,
the widely held belief at the time that living organisms could
arise from nonliving matter. Pasteur showed that if nutrient solu-
tions are freed of all microorganisms (typically by heating) and
protected from airborne contamination, they remain microbe-free
unless and until microorganisms are introduced. Pasteur’s work
on spontaneous generation forced him to develop effective ster-
ilisation procedures, many of which have remained mainstays in
microbiology and clinical medicine to this day.
Pasteur went on from his seminal work on spontaneous gen-
eration to a series of triumphs in medical microbiology. These
included the development of a vaccine against the otherwise
fatal disease rabies and the demonstration that attenuated vac-
cines, made from noninfectious but still active microbes, are safe
and typically more effective than killed vaccines. These were
some of the rst practical successes in the eld of infectious dis-
ease microbiology. However, despite the extensive work of Pas-
teur with various pathogenic agents (see also:Pasteur, Louis),
denitive proof of cause and effect with any infectious disease
remained elusive until the work of Robert Koch.
Robert Koch was a medical doctor primarily interested in
infectious diseases and, in particular, the clear identication
of causative agents of infectious diseases. Koch surmised that
such studies would require the development of methods to
obtain laboratory cultures of suspected disease-causing microbes
(pathogens), and many of the procedures he devised to do this,
such as the use of Petri plates, remain standards in the microbi-
ology laboratory today. From experimental studies on the disease
anthrax and, later, tuberculosis, Koch developed a set of criteria
(known today as Koch’s postulates) that, when faithfully exe-
cuted, unequivocally link a specic microbe to a specic infec-
tious disease.
To full his postulates, Koch and his associates devised meth-
ods to isolate suspected pathogens from diseased animals and
grow them in pure cultures (containing only a single kind of
microbe) in the laboratory. The ability to transmit an infec-
tious disease by injecting the laboratory-cultured pathogen into a
healthy animal was the linchpin in Koch’s postulates and supplied
the denitive proof needed to join cause and effect. In his great-
est medical triumph, Koch used his newly developed laboratory
methods to link the bacterium Mycobacterium tuberculosis with
the disease tuberculosis, and for this monumental achievement,
Koch was awarded a Nobel Prize in 1905. See also:Koch, Hein-
rich Hermann Robert
As microbiology entered the twentieth century, its initial focus
on basic principles, methods and medical aspects broadened to
include studies of the microbial diversity of soil and water and the
metabolic processes that microorganisms carry out in these habi-
tats. Notable microbiologists of this era were Martinus Beijerinck
and Sergei Winogradsky. Beijerinck’s greatest contribution was
his development of the enrichment culture technique, a process
in which highly selective nutrient and incubation conditions are
used to isolate microbes from nature whose metabolism and other
properties are best suited to the conditions employed and thus
give them a competitive advantage. Using this technique, Bei-
jerinck isolated the rst pure cultures of many common soil and
aquatic microorganisms we know today. See also:Beijerinck,
Martinus Willem
Sergei Winogradsky was also interested in the microbial diver-
sity of soils and waters but was particularly interested in the
metabolic reactions carried out by bacteria. Winogradsky was
the rst to show that bacteria can oxidise inorganic nitrogen
and sulphur compounds and that the organisms that oxidised
2eLS © 2015, John Wiley & Sons, Ltd.
nitrogen compounds differed from those that oxidised sulphur
compounds. Further, Winogradsky’s brilliant insight into the
metabolism of these organisms led him to propose the con-
cept of chemolithotrophy, the oxidation of inorganic compounds
for the purpose of obtaining the energy necessary for growth
(Winogradsky, 1949). Winogradsky went on to show that these
organisms – the chemolithotrophs – were widespread in nature
and shared with plants the ability to use CO2as their sole car-
bon source. Winodgradsky was thus the rst to demonstrate that
autotrophy occurred in nonphotosynthetic organisms, a property
that we now know is widespread in the microbial world. See
also:Winogradsky, Sergei Nikolaevitch;Chemolithotrophy;
Microbial Inorganic Carbon Fixation
The modern era of microbiology
The eld of microbiology developed quickly in the twentieth cen-
tury in step with the many new powerful laboratory tools that
became available. During this period, microbiology as a science
matured, spawning several new subdisciplines rooted in genetics
and molecular biology (Table 1). Much of the science of micro-
biology today is fueled by genomics: the mapping, sequencing
and analysis of genes and genomes. New and faster methods of
deoxyribonucleic acid (DNA) sequencing coupled with robust
computer analyses are being used to attack some of the greatest
challenges in medicine, agriculture and the environment, and they
have revealed the true extent and diversity of the microbial world
(López-García and Moreira, 2008). See also:Genome Sequenc-
ing;Genome Mapping
Along with the suite of foundational techniques developed by
early microbiologists, molecular microbiologists today are on
track to understanding how cells work in unprecedented detail.
This knowledge will help humans better exploit the benecial
effects and control the potentially devastating effects of microbial
activities. However, before we consider some of these activities,
we need to compare and contrast the major microbial groups that
compose the microbial world, and we focus on this topic now.
Classification and Basic
Characteristics of Microorganisms
Microorganisms encompass an enormous diversity of micro-
scopic life forms, each with distinct characteristics. On the basis
of their genotypic (genetic) and phenotypic (observed) proper-
ties, all organisms are classied into one of three domains – the
Bacteria,Archaea or Eukarya – and numerous examples of
microorganisms are found in all three (Figure 1;Woeseet al.,
1990). The comparison of ribosomal ribonucleic acid (rRNA)
gene sequences has been especially important in determining
the evolutionary, or phylogenetic, relationships of organisms.
Although very different on a phylogenetic level, the Bacteria
and Archaea (traditionally called prokaryotes) are structurally
similar in that cells of these groups do not typically contain
membrane-bound organelles and, therefore, show a lesser degree
of cellular compartmentalisation than organisms belonging to
the Eukarya (the eukaryotes). The presence of membrane-bound
organelles, including a dened nucleus, is the hallmark of eukary-
otic cells, and microbial Eukarya include fungi, protists and
certain helminths, especially the parasitic worms. An overview
discussion of microbiology must also include a consideration
of viruses, even though they are not cellular and thus are not
included in the three-domain tree of life. We now consider each of
these groups in more detail. See also:Cell Structure;The Cell
Nucleus;Phylogeny Based on 16S rRNA/DNA
The Bacteria and Archaea
The Bacteria and Archaea are vast groups of microorganisms
consisting of potentially hundreds of thousands to millions of
Red algae
Green algae
Bacteria Archaea Eukarya
Crenarchaeota Thaumarchaeota
Figure 1 Universal phylogenetic tree showing relationships between major lineages of the three domains of life (Bacteria,Archaea and Eukarya). Tree
topology and branch lengths were determined by comparative small subunit (SSU) rRNA gene sequence analysis.
eLS © 2015, John Wiley & Sons, Ltd. 3
species, most of which remain uncharacterised. These microbes
are ubiquitous, inhabiting and subsist in nearly every imag-
inable environment on Earth. Various species thrive on or
within every plant and animal, within and underneath mas-
sive glaciers, in hypersaline waters of the Great Salt Lake and
the Dead Sea, and even in boiling hot springs and deep sea
volcanic (hydrothermal) vents. Many microorganisms, called
extremophiles, are able to thrive in environmental conditions
that humans would consider punishingly harsh, and numerous
members of the Archaea, in particular, excel in this lifestyle. See
Bacteria and Archaea are classied based on (1) phylogenetic
distinctions, (2) structural and morphological characteristics,
such as cell shape, size and arrangement and (3) biochemical and
physiological traits, such as growth factor requirements, range of
carbon and energy sources and end products of metabolism. For
example, the Cyanobacteria are a large phylum of physiologi-
cally and phylogenetically related Bacteria that carry out oxy-
genic (O2-producing) photosynthesis using chlorophyll-based
The classication of Bacteria and Archaea into various tax-
onomic groups is rapidly undergoing revision and modica-
tion as novel species continue to be discovered and as insights
from the eld of genomics (a discipline in which the total
genetic makeup of organisms is determined and studied) pro-
vide more accurate information of phylogenetic relationships.
Molecular tools have become essential for classifying Bacte-
ria and Archaea that resist laboratory culture and thus can only
be identied by analysing sequences of DNA or RNA isolated
from natural samples. Once determined, taxonomic conclusions
are published in peer-reviewed journals and organised into ref-
erence manuals, such as Bergey’s Manual of Systematic Bacte-
riology and The Prokaryotes.See also:Semantides and Mod-
ern Bacterial Systematics;Bacterial Cells;Archaeal Cells;
Cells of Bacteria and Archaea exist in three major mor-
phological forms: spherical (coccus), rod-shaped (bacillus) and
spiral-shaped (spirillum) (Figure 2a–c). Less common cell mor-
phologies also exist, such as tightly coiled (spirochete; Figure
2d), appendaged and lamentous. All of these cell types are gen-
erally very small; most rods are 0.51μm wide and 1–4 μm long,
and a typical coccus has a diameter of 0.61μm (about one-tenth
the diameter of a human red blood cell). Many of these cells
are motile by means of one or more rotating appendages called
agella. And unlike animal cells and protozoa, most species
of Bacteria and Archaea have a cell wall, a strong layer of
material located outside the cell membrane that confers struc-
tural integrity and shape to the cell. See also:Bacterial Flag-
ella;Archaeal Flagella;Bacterial Cell Wall;Archaeal Cell
Most species of Bacteria and Archaea divide by binary ssion,
a process in which one cell divides into two identical daughter
cells following replication of the parental cell’s DNA. Bacte-
rial and archaeal genomes are usually organised into a single,
circular chromosome, and many species also have small, extra-
chromosomal, circular pieces of DNA called plasmids, which
often carry genes for antibiotic resistance, toxin production or
specialised metabolisms. The chromosome is densely coiled in
Figure 2 Major morphological forms of bacterial cells. (a) coccus (plural,
cocci); (b) rod (bacillus; plural, bacilli); (c) spirillum (plural, spirilla) and (d)
the cytoplasm to create a nucleoid, which is analogous to the
nucleus of a eukaryotic cell but differs in that it is not bound by
a membrane. See also:Binary Fission in Bacteria;Bacterial
Reproduction and Growth;Bacterial Chromosome;Bacterial
Plasmids;Archaeal Plasmids
Microorganisms of the domain Eukarya
Like Bacteria and Archaea, eukaryotic microorganisms are
remarkably diverse, consisting of hundreds of thousands of
species of fungi, protozoa and algae, as well as hundreds
of species of parasitic worms (Crompton, 1999). Fungi are
a major component of soil ecosystems. Similar to plants,
cells of fungi have rigid cell walls and are nonmotile, but
unlike plants, they lack chlorophyll and are nonphotosynthetic.
Instead, fungi subsist by degrading dead plant and animal mat-
ter, and therefore, along with bacteria, play a key role in the
decomposition and recycling of nutrients. See also:Fungal
Fungi exist in two basic forms: moulds, which consist of la-
ments called hyphae that can form into masses known as mycelia,
and yeasts, which are unicellular and typically oval-shaped. Some
fungi are dimorphic in that they can assume either morphology
depending on environmental conditions. Fungi are capable of
sexual reproduction or asexual reproduction, both of which may
result in the production of spores that can germinate to form new
hyphae. In addition, yeasts often reproduce asexually by budding,
a process in which a new daughter cell develops on the surface of
a parent cell before eventually breaking away. See also:Hyphae;
Fungal Cells;Fungal Spores
Protozoa are unicellular, mostly nonphotosynthetic protists that
lack cell walls. Some protozoa are large enough to be seen with
the unaided eye, although most are microscopic. Many protozoa
are capable of reproducing sexually or asexually. Depending
4eLS © 2015, John Wiley & Sons, Ltd.
on the species, asexual reproduction may occur through any
of several mechanisms, including budding, spore formation or
mitotic ssion. A common mechanism of sexual reproduction in
protozoa is conjugation, in which two cells join, exchange genetic
material and produce progeny by budding or ssion. See also:
Protozoan Asexuality
Protozoa are diverse in their habitats and distribution. Most pro-
tozoa obtain energy by breaking down ingested foods via aerobic
respiration, but some species are anaerobes that lack mitochon-
dria (the energy-generating organelle of eukaryotic cells) and
instead obtain their energy through fermentation. Fermentative
protozoa inhabit anoxic environments, such as the digestive tract
of certain animals, where they often establish symbiotic rela-
tionships that may be either benecial or harmful to their host.
Some protozoa cause devastating human diseases, such as malaria
(caused by species of Plasmodium), while others (e.g. Para m e -
cium) are innocuous members of the biosphere, where they exist
as important components of the food chain. See also:Protozoan
Ecology;Protozoan Symbioses
Algae are plant-like protists that are distinguished from fungi
and most protozoa by their ability to perform photosynthesis
using chlorophyll pigments, and they comprise much of the basis
of the food chain in marine and freshwater environments. Algae
exhibit a variety of morphological forms, including unicellular,
lamentous, colonial and large multicellular aggregates called
kelps or ‘seaweeds’ that can attain lengths of up to 50 m. Some
algae have become increasingly important sources of food or
food additives for humans. For example, the red alga Porphyra,
known as nori, is popular in sushi preparation, and other red algae
are the source of agar, a polysaccharide used as a solidifying
agent to make Petri plate culture media as or as a thickener for a
variety of foods. Most algae reproduce asexually, whereas others
form spores or reproduce by fragmentation of cells from larger
aggregates. Some algae reproduce sexually by forming diploid
zygotes from haploid gametes. See also:Algal Ecology;Algal
The helminths are a group of multicellular animals that includes
roundworms and atworms. Even though some of these organ-
isms are visible to the naked eye, they are important topics of
study for microbiologists because (1) they often have microscopic
larval forms as part of their life cycles and (2) many species of
helminths are parasitic and cause important infectious diseases.
Helminth life cycles can be complex and often require multiple
hosts for the different stages. In addition to humans, these hosts
may include other mammals, insects (e.g. ies and mosquitoes),
various sh species and certain aquatic invertebrates, such as
squid, snails and crustaceans.
A helminth infection usually begins with either an insect bite
or an accidental ingestion of worm eggs or larvae. However, a
few worm species, such as those that cause hookworm and schis-
tosomiasis, are capable of burrowing directly through the skin.
The most important means of preventing parasitic worm infec-
tions include thoroughly cooking foods, drinking only puried
or boiled water and employing repellents or physical barriers
to prevent insect bites. Effective treatment of established infec-
tions is often possible using antihelminthic drugs. See also:
Schistosomiasis and Other Trematode Infections
Viruses and virology
Viruses are acellular microbes that require living host cells to
multiply; thus, they are obligate intracellular parasites. Struc-
turally, viruses are quite simple, often consisting of only DNA
or RNA (the viral genome) surrounded by a simple protein coat
having either a helical or icosahedral morphology (Figure 3).
Viruses that infect bacteria, called bacteriophages, often have a
complex morphology in that they exhibit a combination of these
two forms (Figure 3). Most viruses are too small to be seen with
even the best light microscopes, and because of their tiny size
and dependence on host cells, their genomes are typically quite
small, in some cases consisting of only two genes (Faurez et al.,
2009; Niagro et al.,1998)! See also:Viru ses;Virus Structure;
Viruses replicate within an infected cell by commandeering
the host’s enzymatic machinery, which may include use of the
host cell’s nucleic acid polymerases (enzymes that make DNA
or RNA) and/or ribosomes. Following viral replication, progeny
viruses are released, either by lysis of the host cell or by bud-
ding from the host cell’s membrane. Therefore, in many cases,
viral infections lead to death of the host cell, either abruptly
or eventually. Viruses cause many serious diseases, includ-
ing acquired immunodeciency syndrome (AIDS), inuenza,
measles, poliomyelitis, rabies and haemorrhagic fevers, such as
Ebola. See also:Virus Replication
With a broad overview of microbial diversity in place, we can
now explore the crucial role that microorganisms play in the
environment around us.
Microbial Ecology
Microbial ecology is focused on how microbial communities
interact with each other and their environments. A microbial
community is an assembly of one or more populations of cells,
each population composed of a single kind or species of microbe.
An ecosystem is a dynamic complex of living organisms and their
abiotic surroundings, all of which interact as a functional unit.
In terrestrial and aquatic ecosystems, microorganisms interact
with each other and with the plants and animals in the ecosystem.
Microbes play essential roles in these ecosystems by cycling
inorganic nutrients and both producing and consuming organic
matter. Associations of specic microbes with specic plants or
animals are also quite common, and many of these associations
are essential for the health and well-being of the plant or animal.
We briey consider the roles of microorganisms in some major
ecosystems now.
Soil, water and higher organisms as
homes for microbial communities
Soil is the loose outer material that comprises much of Earth’s
surface and forms over long periods of time from a combination
of biological and chemical processes. Soils often contain large
numbers of microorganisms, and depending on the amount of
organic matter present and the soil pH, salinity, degree of aeration
and other abiotic factors, soil microbial communities can be
eLS © 2015, John Wiley & Sons, Ltd. 5
Ebola virus
500 nm
Escherichia coli
Figure 3 Major morphological forms of viruses. Most viruses are considerably smaller than cells (a representation of an Escherichia coli cell is shown for
size comparison). Viruses of eukaryotes typically have either icosahedral (e.g. rhinoviruses) or helical (e.g. ebola virus) symmetry, whereas viruses that infect
bacteria (bacteriophages) often exhibit a complex ‘head-and-tail’ combination of these forms.
relatively simple or highly complex. In addition to soil, however,
microbial communities also exist deep underground in the Earth’s
subsurface, fed by nutrients transported by groundwater.
In soils, microbial numbers are typically greatest in and around
plant roots, a zone called the rhizosphere. It is here that organic
matter excreted from the roots and from dead plant material
greatly stimulates the activities of microbial communities. Tem-
poral changes in the abundance and composition of soil microbial
communities occur from variations in moisture, organic matter
inputs and temperature. In contrast to surface soil, subsurface
microbial communities are less dynamic due to more predictable
conditions. Various Bacteria,Archaea, microbial eukaryotes and
viruses inhabit soils and the deep subsurface, and many important
nutrient cycling reactions occur there, including major transfor-
mations of the elements C, N and S, key constituents of living
Aerobic bacteria and fungi in soils consume oxygen in their
respiratory activities. Many bacteria are anaerobes, carrying out
various types of fermentation or anaerobic respiration (a form of
respiration in which an oxidant other than O2is used). However,
in the nal analysis, aerobic and anaerobic respiration and fer-
mentation all accomplish the same thing; these metabolisms oxi-
dise organic carbon, returning it to CO2. Many anoxic (O2-free)
zones exist in soils and thus aerobes and anaerobes coexist there.
Aquatic microbial ecosystems include both freshwaters and
ocean waters. These two environments differ in many ways,
including salinity, average temperature, depth and nutrient levels.
Photosynthetic microbial communities play important roles in
aquatic ecosystems. The CO2-xing (autotrophic) activities of
these organisms provide not only the organic carbon needed for
their own metabolism and growth but also the organic matter and
oxygen needed by heterotrophic microbes present in the aquatic
microbial community. Photosynthetic microbes are indeed the
base of the aquatic food chain and thus are critical components
of aquatic ecosystems.
The sediments of fresh and marine waters are hotbeds of
distinctly different anaerobic metabolisms. In freshwater sedi-
ments, the bulk of organic carbon is eventually degraded to CH4
(methane). Methane is formed by methanogenic Archaea that
reduce CO2to CH4using hydrogen (H2) as reductant in the
anaerobic respiration called methanogenesis. In contrast to fresh-
water sediments, marine sediments contain large amounts of salts,
including sulphate (SO4
2). Sulphate respiration, whereby SO4
is reduced to H2S, is the dominant form of anaerobic respiration
in marine sediments because it is more energetically favourable
than methanogenesis (the low levels of SO4
2in most freshwater
sediments limits sulphate reduction and favours methanogenesis)
(Widdel and Bak, 1992).
Many microorganisms form relationships with other organ-
isms, which can include other microbes, plants or animals. Such
associations are called symbioses (literally, ‘living together’).
Plants interact with microorganisms through their roots and leaf
surfaces, but in some cases, the association becomes highly spe-
cic and intimate, including actual growth within the plant tis-
sue. Such is the case with the legume–root nodule symbiosis,
an association in which tumour-like nodules form on the roots
of leguminous plants (plants that bear their seeds in pods), such
as soybeans and peanuts. The root nodules provide a habitat for
6eLS © 2015, John Wiley & Sons, Ltd.
bacteria that x atmospheric nitrogen into ammonia (N2+6H
2NH3). The plant then uses the ammonia as a source of nitro-
gen to make proteins and nucleic acids, allowing it to thrive in
nitrogen-poor soils.
Animal–microbe symbioses are also quite common. The
human large intestine, for example, contains enormous numbers
of bacterial cells that form a huge microbial community, the
human gut microbiome, the composition of which varies from
person to person and is inuenced by diet, health and other
factors (The Human Microbiome Project Consortium, 2012). As
more is learned about the microbiomes of different animals, it
has become clear that any perturbations in this species-specic
microbial community can affect an animal’s physical condition
and susceptibility to disease. In some cases, the microbiome is
essential for the very nutrition of the animal. For example, in
the rumen (forestomach) of animals such as cows and sheep,
bacteria and protozoa digest cellulose and ferment the released
glucose to fatty acids, which are taken up by the animal. This
allows the ruminant to subsist on a diet of plant matter, which is
primarily cellulose.
Microbial nutrient cycling
The key nutrients for life are cycled by both microorganisms
and by plants and animals, but for any given nutrient, it is
microbial activities that dominate. The major nutrient cycles
include those of carbon, nitrogen and sulphur (Figure 4). Car-
bon is cycled primarily through CO2(carbon dioxide) and the
large pool of organic compounds present in living organisms
(Figure 4a). CO2is reduced to organic compounds by plants
but also by many different microorganisms, both photosynthetic
and chemolithotrophic. Organic matter, either excreted by living
organisms or released from dead organisms, is eventually oxi-
dised to CO2or is converted to methane (CH4) by methanogenic
archaea and later oxidised to CO2by methane-consuming bacte-
ria. See also:Global Carbon Cycle
Nitrogen compounds can be either oxidised or reduced by
microbes depending on the compound. Ammonia (NH3) is oxi-
) by a group of chemolithotrophs called
the nitrifying bacteria. Nitrate is then reduced to atmospheric
nitrogen (N2) by nitrate-respiring anaerobic bacteria (denitri-
ers) or to NH3by ammonifying bacteria. The major remaining
link in the nitrogen cycle is the reduction of N2to NH3by the
nitrogen-xing bacteria (Figure 4b). This process is an impor-
tant means of enriching soils in usable nitrogen and is the key to
the soybean–root nodule symbiosis mentioned earlier. See also:
Nitrogen Fixation;Nitrication
Sulphur is cycled primarily between SO4
2and sulphide (H2S)
through the activities of sulphate-reducing (sulphate-respiring)
bacteria and chemolithotrophic sulphide-oxidising bacteria. Ele-
mental sulphur (S0) is often an intermediate product in these
metabolisms and can be either oxidised to sulphate or reduced
to sulphide (Figure 4c). Sulphur-oxidising bacteria are primarily
aerobes and participate in the sulphur cycle by consuming sul-
phide – a toxic substance for higher organisms – and generating
sulphate, a key plant nutrient. By contrast, sulphate-reducing bac-
teria are anaerobes and oxidise organic matter to CO2in anoxic
environments, generating sulphide (Figure 4c). Some photosyn-
thetic bacteria can also oxidise sulphide to support autotrophic
growth (CO2+H2SOrganic matter +SO4
2)(Gregersenet al.,
2011). Other key elements, such as P and Fe, are also cycled in
nature, and like the C, N and S cycles, the cycling of these nutri-
ents is driven primarily by the activities of microorganisms. See
also:Sulfur Oxidation in Prokaryotes;Biogeochemical Cycles
Medical Microbiology
While the vast majority of microorganisms provide benecial
services to humans, a few species (<1%), called pathogens, are
potentially harmful. When pathogens successfully invade, mul-
tiply and cause damage to the host, disease ensues. Pathogens
can damage the host in two ways: toxicity and invasiveness. Sev-
eral pathogens produce toxins that can harm or even be lethal to
humans. Some toxins, called exotoxins, are proteins secreted by
the pathogen once established in the host. By contrast, endotox-
ins make up part of the outer layer (cell wall) of the cell itself.
Bacterial exotoxins are among the most potent toxins known.
Carbon Nitrogen Sulphur
N2, N2O, NO
(a) (b) (c)
Figure 4 Key elemental cycles driven predominantly by the activity of microorganisms. (a) Carbon is cycled between inorganic (CO2) and organic (CH2O)
forms by the actions of autotrophs (1) and heterotrophs (2). In addition, methane (CH4) is produced (3) or consumed (4) by methanogens or methanotrophs,
respectively. (b) Nitrogen compounds are cycled by nitrogen-fixing (1), nitrosifying (2), nitrifying (3) and denitrifying (4) bacteria. (c) In the sulphur cycle,
sulphide is produced by sulphate- and sulphur-reducing bacteria (1 and 2, respectively). Sulphur chemolithotrophic bacteria oxidise sulphide to sulphate
through an elemental sulphur intermediate (3).
eLS © 2015, John Wiley & Sons, Ltd. 7
For example, just 1 μg (one-millionth of a gram) of botulinum
exotoxin can kill an adult human (Arnon et al., 2001). See also:
Botulinum Toxin;Toxin Action: Molecular Mechanisms
Invasive pathogens harm the host by directly damaging tis-
sues or depriving them of nutrients. Bacterial invasion begins
with attachment to host cells, which is facilitated by cellular
appendages, such as mbriae or pili. Tissue destruction occurs
through the secretion of enzymes, such as leukocidins, which
destroy white blood cells, and collagenases, which erode colla-
gen, a brous protein found in various connective tissues. Some
bacteria produce dense polymer coatings outside their cell walls
called capsules that protect the invading pathogen from the host’s
immune response. Hosts respond to microbial invasion and the
presence of toxins by producing protective antibodies that bind to
and neutralise the microbe or toxin and by amassing white blood
cells to attack invading pathogens by either consuming them
or producing toxic agents to kill them. See also:Bacterial Pili
and Fimbriae;Bacterial Capsules and Evasion of Immune
Human hosts acquire infectious diseases by different routes,
including the respiratory tract, the oral cavity and digestive
system and the skin and genitourinary system. In some cases,
pathogens are transmitted person-to-person, as in sexually trans-
mitted infections (e.g. gonorrhoea and syphilis) or respiratory
diseases (e.g. tuberculosis and inuenza). In other instances,
pathogens are transmitted indirectly through inanimate objects,
such as clothing or towels, or through vectors, animals that
carry pathogens between susceptible hosts. Lyme disease and
malaria are examples of diseases that are transmitted indirectly
to humans by arthropod vectors, the deer tick and Anopheles
mosquito, respectively. After a pathogen is transmitted to a sus-
ceptible host, an incubation period occurs in which the pathogen
multiplies and becomes established in the host before signs and
symptoms of disease appear. See also:Syphilis: Epidemiolog-
ical Aspects;Tuberculosis;Malaria;Respiratory System:
Bacterial Infections
Antibiotics are antimicrobial chemicals produced by various
microorganisms. Antibiotics are often administered to help ght
infectious microorganisms and kill or inhibit pathogens by dis-
rupting nucleic acid or protein synthesis, damaging the plasma
membrane, preventing cell wall synthesis or interfering with cel-
lular metabolism. Viruses, which replicate only inside a host cell,
are not affected by antibiotics and must therefore be treated by
other chemical compounds that block viral enzymes or alter viral
nucleic acids. The indiscriminate use of antibiotics in humans
and other animals in recent years has led to the development of
microorganisms that are resistant to many of these drugs. These
antibiotic-resistant microbes challenge conventional chemother-
apy and are of major concern to medical professionals.
An alternative strategy to combating microbial infections is to
employ vaccines to prevent pathogens from becoming established
in the body in the rst place. Vaccines induce an immune response
in the host, and this triggers the production of protective antibod-
ies that provide immunity against specic infectious agents and
other foreign antigens. Vaccines have been particularly helpful
in preventing common childhood diseases, including diphtheria,
whooping cough (pertussis), poliomyelitis, measles and mumps.
See also:Antiviral Drugs;Antibiotics and the Evolution of
Antibiotic Resistance;Vaccination of Humans
Applications in Microbiology
Commercial products from
Microorganisms can be harnessed to make many valuable prod-
ucts, and industrial microbiology and biotechnology are the sub-
disciplines of microbiology focused on these tasks (Table 1).
Industrial microbiology uses microorganisms to synthesise prod-
ucts in large amounts. This is done by taking microbes that
naturally produce some substance of relatively low value – for
example, an antibiotic or alcohol – and selecting for ‘overpro-
ducing strains’ that can be grown on a huge scale; the resulting
product may be made by tons or thousands of litres. Biotech-
nology, by contrast, employs genetically engineered microbes to
synthesise small amounts of very high-value products that the
microbes are otherwise unable to make, such as a human protein.
Some major products of industrial microbiology and biotechnol-
ogy are listed in Table 2.
Food and mining microbiology
Food production and mining are obviously quite different activ-
ities but both owe their success to the microbial world. Certain
microbes are used in the preparation of common food products
while others are used in the mining industry to extract valuable
minerals from crude ores.
Yeast is the key catalyst in the production of baked goods and
alcoholic beverages. The fermentative metabolism of yeasts (glu-
cose 2 ethanol +2CO
2) generates key products for the baker
(CO2to raise the dough) and the brewer (ethyl alcohol). Many
cheeses owe their characteristic avours to the activities of fer-
mentative microbes. For example, the key components in Swiss
Table 2 Selected products of industrial microbiology and
Product Example
Industrial microbiology
Antibiotics Penicillin, tetracycline
Enzymes Laundry proteases and lipases,
glucose isomerase
Food additives Vitamins, amino acids, citric acid
Alcohol/chemicals Ethanol in alcoholic beverages and
gasohol, butanol, biodiesel and
Human hormones Insulin, somatotropin
Blood proteins Tissue plasminogen activator
Immune modulators Interferons, tumour necrosis factor
Therapeutic enzymes DNase
8eLS © 2015, John Wiley & Sons, Ltd.
cheese – propionic acid and CO2– are responsible for the dis-
tinctive nutty taste and characteristic holes (eyes), respectively, of
this popular cheese. Other major food products of microbial ori-
gin include fermented milks (e.g. buttermilk and yogurt), meats
(e.g. salami and summer sausage) and vegetables (e.g. sauerkraut
and pickles).
The mining of copper, gold and a few other metals relies on
the leaching properties of bacteria. These valuable metals are
typically present in low amounts in complex mineral ores that
contain large amounts of iron. The metabolic activities of various
acid-tolerant, iron-oxidising bacteria release these metals from
the ores and generate acidic solutions (the leachate) from which
the now solubilised metals can be concentrated and harvested by
chemical treatment. Copper, widely used in electrical equipment,
piping and the brewing industry, is the most commonly leached
metal. The strategic element uranium is also mined by microbial
leaching processes.
Environmental applications of
Microorganisms play critically important roles in the environ-
ment. We have already considered the key roles that microbes
play in nature’s major nutrient cycles. But in addition to these
more or less continuous activities, microorganisms have been
exploited for purifying wastewaters and for cleaning pollution in
the environment, a process called bioremediation.
Sewage and other wastewaters must be treated before they can
be released into natural waterways. This is because without treat-
ment, the massive inux of organic matter and mineral nutrients
would trigger extensive microbial growth and O2consumption,
causing die-offs of plants and animals and diminishing the aes-
thetic and recreational value of the water. To deal with the high
nutrient load of wastewaters, elaborate treatment facilities are
used to stimulate the activities of complex microbial communities
to remove as much organic carbon and other polluting nutrients
(such as nitrates and phosphates) from the wastewaters as pos-
sible. Following treatment, the water can then be safely released
into rivers or other bodies of water. See also:Eutrophication of
Lakes and Rivers
To produce potable drinking water, additional treatment is
necessary to remove as many potentially pathogenic microor-
ganisms and remaining toxic substances as possible. Drinking
water production includes the coagulation and ltration of already
high-quality surface or subsurface waters followed by disinfec-
tion with chlorine and transport of the water through water mains
to the consumer. The entire process of drinking water production
must be carefully performed and monitored to prevent break-
downs that can lead to incidents of serious waterborne illness,
such as cholera or typhoid fever.
When pollution of the environment occurs, either from natural
events or from the activities of humans, microorganisms can be
harnessed to clean up the mess. Microbial bioremediation is typ-
ically the most cost-effective method of removing environmental
pollutants and, in many cases, it is the only practical way to
accomplish the job. Bioremediation is grounded in the astounding
diversity of metabolic reactions capable in the microbial world.
Thus, if some pollutant, such as crude oil, is spilled in the environ-
ment, oil-consuming microbes applied to the spill site can clean
up the mess by oxidising hydrocarbons in the oil to CO2.See also:
In a similar manner, microbes that can degrade pesticides, such
as insecticides and herbicides, are benecial in keeping these
poisonous substances from accumulating in the environment and
damaging plants and animals that were not the original targets
of these agents. Although not every substance that humans have
created is biodegradable (e.g. teon is not), the vast majority of
pollutants are, and it is through the activities of microorganisms
that these undesirable substances are converted into compounds
that can enter the natural nutrient cycles (Figure 4).
Humans owe a considerable debt to the microbial world for
keeping planet Earth habitable and healthy. If cyanobacteria had
never become established on Earth, then the oxygen we breathe
and depend on would never have been produced. And if it were
not for microbes today, the everyday activities of humans would
eventually damage the environment beyond its capacity to sustain
human life. The microbial world is clearly the foundation of the
biosphere, and thus the science of microbiology, which attempts
to understand this unusual world, may be our most relevant
biological science today.
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Further Reading
Brock TD (1961) Milestones in Microbiology. Upper Saddle River,
NJ: Prentice Hall.
Dixon B (2009) Animalcules: The Activities, Impacts, and Investiga-
tors of Microbes. Washington, DC: ASM Press.
Henig RM (1993) A Dancing Matrix: Voyages Along the Viral Fron-
tier. New York, NY: Alfred A. Knopf, Inc.
Kirchman DL (2012) Processes in Microbial Ecology.NewYork,
NY: Oxford University Press, Inc..
Kohanski MA, Dwyer DJ and Collins JJ (2010) How antibiotics kill
bacteria: from targets to networks. Nature Reviews Microbiology
8: 423–435.
Konstantinidis KT and Tiedje JM (2007) Prokaryotic taxonomy
and phylogeny in the genomic era: advancements and challenges
ahead. Current Opinions in Microbiology 10: 504–509.
Strelkauskas AJ, Strelkauskas JE and Moszyk-Strelkauskas D (2010)
Microbiology: A Clinical Approach. New York, NY: Garland Sci-
ence, Taylor and Francis Group, LLC.
Tortora GJ, Funke BR and Case CL (2016) Microbiology: An
Introduction, 12th edn. San Francisco, CA: Benjamin Cum-
10 eLS © 2015, John Wiley & Sons, Ltd.
ResearchGate has not been able to resolve any citations for this publication.
Full-text available
Objective The Working Group on Civilian Biodefense has developed consensus-based recommendations for measures to be taken by medical and public health professionals if botulinum toxin is used as a biological weapon against a civilian population.Participants The working group included 23 representatives from academic, government, and private institutions with expertise in public health, emergency management, and clinical medicine.Evidence The primary authors (S.S.A. and R.S.) searched OLDMEDLINE and MEDLINE (1960–March 1999) and their professional collections for literature concerning use of botulinum toxin as a bioweapon. The literature was reviewed, and opinions were sought from the working group and other experts on diagnosis and management of botulism. Additional MEDLINE searches were conducted through April 2000 during the review and revisions of the consensus statement.Consensus Process The first draft of the working group's consensus statement was a synthesis of information obtained in the formal evidence-gathering process. The working group convened to review the first draft in May 1999. Working group members reviewed subsequent drafts and suggested additional revisions. The final statement incorporates all relevant evidence obtained in the literature search in conjunction with final consensus recommendations supported by all working group members.Conclusions An aerosolized or foodborne botulinum toxin weapon would cause acute symmetric, descending flaccid paralysis with prominent bulbar palsies such as diplopia, dysarthria, dysphonia, and dysphagia that would typically present 12 to 72 hours after exposure. Effective response to a deliberate release of botulinum toxin will depend on timely clinical diagnosis, case reporting, and epidemiological investigation. Persons potentially exposed to botulinum toxin should be closely observed, and those with signs of botulism require prompt treatment with antitoxin and supportive care that may include assisted ventilation for weeks or months. Treatment with antitoxin should not be delayed for microbiological testing.
Full-text available
Green sulfur bacteria (GSB) constitute a closely related group of photoautotrophic and thiotrophic bacteria with limited phenotypic variation. They typically oxidize sulfide and thiosulfate to sulfate with sulfur globules as an intermediate. Based on genome sequence information from 15 strains, the distribution and phylogeny of enzymes involved in their oxidative sulfur metabolism was investigated. At least one homolog of sulfide:quinone oxidoreductase (SQR) is present in all strains. In all sulfur-oxidizing GSB strains except the earliest diverging Chloroherpeton thalassium, the sulfide oxidation product is further oxidized to sulfite by the dissimilatory sulfite reductase (DSR) system. This system consists of components horizontally acquired partly from sulfide-oxidizing and partly from sulfate-reducing bacteria. Depending on the strain, the sulfite is probably oxidized to sulfate by one of two different mechanisms that have different evolutionary origins: adenosine-5'-phosphosulfate reductase or polysulfide reductase-like complex 3. Thiosulfate utilization by the SOX system in GSB has apparently been acquired horizontally from Proteobacteria. SoxCD does not occur in GSB, and its function in sulfate formation in other bacteria has been replaced by the DSR system in GSB. Sequence analyses suggested that the conserved soxJXYZAKBW gene cluster was horizontally acquired by Chlorobium phaeovibrioides DSM 265 from the Chlorobaculum lineage and that this acquisition was mediated by a mobile genetic element. Thus, the last common ancestor of currently known GSB was probably photoautotrophic, hydrogenotrophic, and contained SQR but not DSR or SOX. In addition, the predominance of the Chlorobium-Chlorobaculum-Prosthecochloris lineage among cultured GSB could be due to the horizontally acquired DSR and SOX systems. Finally, based upon structural, biochemical, and phylogenetic analyses, a uniform nomenclature is suggested for sqr genes in prokaryotes.
The recent discovery of numerous detrital submicron particles in diverse marine environments (Koike et al., 1990; Longhurst et al., 1992; Wells and Goldberg, 1991, 1994) has stirred the interest of oceanographers and has spurred studies into the roles of these small particles in marine food webs and biogeochemical fluxes. The abundance of non-living submicron particles (107–1010 particles ml−1) far exceeds the number of living particles of similar size dimensions, including phytoplankton, bacteria, and viruses (Koike et al., 1990; Wells and Goldberg, 1991; see Table I). Bulk chemical measurements have confirmed that the “colloidal fraction” (size, 0.001–1 μm) represents a large fraction (10–50%) of total “dissolved” organic carbon (DOC) in seawater (Ogawa and Ogura, 1992; Benner et al., 1992; Gau et al., 1994). Several provocative hypotheses have been proposed to explain the roles of colloids and submicron particles in trophic dynamics (Sherr, 1988; Flood et al., 1992), aggregate formation (Alldredge et al., 1993; Kepkay, 1994), and condensation of organic matter (Nagata and Kirchman, 1992b, 1996; Keil and Kirchman, 1994).
An overview of the sulfate-reduction process is given in Chapter 24. Most types of dissimilatory sulfate-reducing bacteria that have been isolated from nature and described so far are mesophilic, nonsporeforming anaerobes. They are members of the delta subdivision of the proteobacteria. The earliest known representative of this category is Desulfovibrio (Beijerinck, 1895). Further investigations have revealed a great morphological and nutritional diversity within this group. Various cell types have been described including cocci; oval or long straight rods; more or less curved rods or spirilla; cell packets; cells with gas vesicles; and gliding, multicellular filaments (Figs. 7–9). Electron donors used for sulfate reduction include H2, alcohols, fatty acids, other monocarboxylic acids, dicarboxylic acids, some amino acids, a few sugars, phenyl-substituted acids, and some other aromatic compounds (Table 2). Even long-chain alkanes can be anaerobically oxidized by a particular type of sulfate-reducing bacterium (Aeckersberg et al., 1991). The utilization of polysaccharides or polypeptides, such as has been observed with the extremely thermophilic sulfate-reducing archaebacterium Archaeoglobus (Stetter, 1988; Stetter et al., 1987), has not been reported for mesophilic sulfate reducers.