S-BIOL316 GENERAL MICROBIOLOGY LECTURE

MODULE 3.0

MICROBIAL TAXONOMY
AND SYSTEMATICS

https://en.wikipediam.org/wiki/Three-domain_system

Biological Sciences Department

College of Science and Computer Studies
De La Salle University-Dasmariñas
Dasmariñas City, Cavite

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I. INTRODUCTION

Taxonomy and systematics are two concepts related to the study of diversification of
living forms and the relationships of living things through time. Taxonomy involves not just
naming organisms but grouping them with other organisms that share common properties.
The main difference between taxonomy and systematics is that taxonomy is involved in the
classification and naming of organisms whereas systematics is involved in the
determination of evolutionary relationships of organisms. This means systematics ascertain
the sharing of the common ancestry by different organisms.

In taxonomy, different organisms are scientifically named and grouped in different

taxonomic levels. In systematics, organisms are grouped based on their evolutionary
relationships. Taxonomy can be considered as a branch of systematics. Both taxonomy
and systematics use morphological, behavioral, genetics, and the biochemical
observations. For more reading on the difference, refer to this url:
https://www.researchgate.net/publication/320413728_Difference_Between_Taxonomy_and
_Systematics and Madigan et al. 2013.

II. HISTORY OF CLASSIFICATION SYSTEM

In the early days, classification appeared relatively straightforward, with all living things
apparently fitting into one of two kingdoms of plants and animals. Most of us are
accustomed to the Linnaean system of classification that assigns every organism a
kingdom, phylum, class, order, family, genus, and species, which, among other
possibilities, has the handy mnemonic King Philip Came Over For Good Soup. It was in
1735 when the two-kingdom system, Plantae and Animalia was formally introduced by
Swedish botanist Carolus Linnaeus. This system was created long before scientists
understood that organisms evolved.

The inclusion of microorganism in the Linnean system became apparent in 1857

when Carl von Nägeli, proposed to include bacteria and fungi in the plant kingdom. In 1866,
Ernst Haeckel proposed the Kingdom Protista, to include bacteria, protozoa, algae, and
fungi. The term prokaryote was then introduced in 1937 by Edouard Chatton to distinguish
cells having no nucleus from the nucleated cells of plants and animals, with the current
definition provided by Roger Stanier in 1961 - “cells in which the nuclear material
(nucleoplasm) is not surrounded by a nuclear membrane”.

In 1968, Robert G.E. Murray proposed the Kingdom Prokaryotae. In 1969, Robert H.
Whittaker founded the five-kingdom system in which prokaryotes were placed in the
Kingdom Prokaryotae, or Monera, and eukaryotes comprised the other four kingdoms. The
Kingdom Prokaryotae had been based on microscopic observations. Subsequently, new
techniques in molecular biology revealed that there are actually two types of prokaryotic
cells and one type of eukaryotic cell. Increased focus on the cellular and molecular

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similarities and dissimilarities between organisms led to proposals for further refinements to
the five-kingdom system proposed by Robert Whittaker in 1969.

Because the Linnaean system is not based on evolution, most biologists are
switching to a classification system that reflects the organisms' evolutionary history.
Phylogenetic classification system names only clades — groups of organisms that
descended from a common ancestor. As an example, we can look more closely at reptiles
and birds. Read more about this system in this URL:
https://evolution.berkeley.edu/evolibrary/article/evo_10

III. PHYLOGENETIC TREE OF LIFE

DNA sequence-based phylogenetic analysis has revealed that the five kingdoms do
not represent five primary evolutionary lines. Instead, cellular life on Earth has evolved
along three primary lineages, called domains (Figure 1). Two of these domains, the
Bacteria and the Archaea, are exclusively composed of prokaryotic cells. The Eukarya
contains the eukaryotes including the plants, animals, fungi, and protists.

IV.
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Figure 1. Phylogenetic Relationships of the Prokaryotes based on 16S rRNA. (Madigan et
al. 2013)

Molecular studies in the 1970s revealed that the Archaea differed from all other bacteria
in their 16S rRNA sequences, as well as in their cell wall structure, membrane lipids and
aspects of protein synthesis. These differences were seen as sufficiently important for the
recognition of a third basic cell type to add to the prokaryotes and eukaryotes. This led to the

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proposal of a three-domain scheme of classification, in which prokaryotes are divided into the
Archaea and the Bacteria.

Ribosomal RNAs (rRNA), are excellent tools for discerning evolutionary relationships
because all cells contain ribosomes (and thus rRNA), including microorganisms. Carl Woese,
an American microbiologist, pioneered the use of comparative rRNA sequence analysis as a
measure of microbial phylogeny and, in so doing, revolutionized our understanding of cellular
evolution. In brief, genes encoding rRNA from two or more organisms are sequenced and the
sequences aligned and scored, base-by-base, for sequence differences and identities using a
computer; the greater the sequence variation between any two organisms, the greater their
evolutionary divergence. Then, using a treeing algorithm, this divergence is depicted in the
form of a phylogenetic tree.

The gene for 16S rRNA which are highly conserved and highly variable regions was
used to determine the three lineages. Based on the analysis, three cell lineages clearly
emerged as cells were forming 3.5 billion years ago. This led to the Archaea, the Bacteria, and
what eventually became the nucleoplasm of the eukaryotes. The phylogenetic tree of life
reveals two very important evolutionary facts: (1) all prokaryotes are not phylogenetically
closely related, and (2) Archaea are actually more closely related to Eukarya than to Bacteria.
Thus, from the last universal common ancestor (LUCA) of all life forms on Earth, evolutionary
diversification diverged to yield the ancestors of the Bacteria and of a second main lineage.
The latter once again diverged to yield the ancestors of the Archaea, a lineage that retained a
prokaryotic cell structure, and the Eukarya, which did not. The universal tree of life shows that
LUCA resides very early within the Bacteria domain.

3.1 Domain Bacteria

In the Domain Bacteria, there are already 80 lineages discovered. However, only the
key ones are shown in such a way that their phenotypic traits are well known and studied since
they are already cultured in the laboratory. Although some lineages are characterized by
unique phenotypic traits, such as the morphology (the cell shape of spirochetes, physiology of
cyanobacteria) most species of bacteria were classified by molecular analysis since these
bacteria lack phenotypic cohesiveness, or they share almost the same traits as the other
species. The largest group, the Proteobacteria, is a good example of this, as collectively this
group shows all known forms of microbial physiology.

As Figure 2 shows, the most phylogenetically ancient phylum contains the genus
Aquifex, which are hyperthermophilic H2 (Hydrogen) – oxidizing chemolitotrophs. These were
followed by the Thermodesulfobacterium, Thermotoga, and the Chloroflexus group (green non-
sulfur bacteria) which do also contain thermophilic species. Furthermore after the green non-
sulfur bacteria, the Deinococci, the unique Spirochetes, the phototrophic green sulfur bacteria,
the chemoorganotrophic Flavobacterium and Cytophaga groups, the budding Planctomyces
and the Verrucomicrobium groups, the Chlamydia, and the genera Nitrospira and
Deferribacter. Other major groups include the gram-positive bacteria and the Cyanobacteria.
The Firmicutes and the Actinobacteria, which are the Gram-positive bacteria, come after.

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Figure 2. Phyla of Domain Bacteria (Madigan et al. 2013)

3.2 Domain Archaea

Carl Woese, along with his colleagues, constructed the phylogenetic trees for the
prokaryotes, which showed evolutionary relatedness. Woese’s work also revealed that one
group of prokaryotes are very different from all the others. This group of prokaryotes are
known as Archaea, which differ from bacteria by its cell wall and plasma membrane chemistry,
as well as the sequences of its 16S rRNA. Archaea share some features which are similar
with other bacteria and some with Eukaryotes in Table 1.

Archaean members show diversity of both morphological and physiological

characteristics. However, it was not noticed before since Archaea remained unidentified as a
separate group because they did not display any very obvious morphological differences from
true bacteria like the main cell shapes for example. This is a problem before since one of the
first ways of classifying bacteria are by morphological characteristics.

Archaea Bacteria Eukarya

Characteristics
Main genetic Single closed circle of Single closed circle True nucleus with

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material dsDNA of dsDNA multiple linear

chromosomes
Histones Present Absent Present
Gene structure Introns absent Introns absent Introns present
Plasmids Common Common Rare
Polycistronic Present Present Absent
mRNA
Ribosomes 70S 70S 80S
Protein Synthesis Not sensitive to Sensitive to Not sensitive to
streptomycin, streptomycin, streptomycin,
chloramphenicol chloramphenicol chloramphenicol
Initiator tRNA Methionine N-formyl methionine Methionine
Membrane fatty Ether-linked, branched Ester-linked, straight Ester-linked straight
acids chain chain
Internal organelles Absent Absent Present
Site of energy Cytoplasmic Cytoplasmic Mitochondria
generation membrane membrane
Cell wall Muramic acid absent Muramic acid present Muramic acid absent

Table 1. Similarities and Difference of the Characteristics of the Three Domains of Life.

However, as more advances in microbial research push through, more unusual shapes
of Archaean members are encountered, members of the genus Haloarcula for example, have
flattened square or triangular cells (Figure 3). Gram-positive and Gram-negative forms of these
members are also found. However, neither group possesses a true peptidoglycan. Some
members of the said domain possess pseudomurein, which is composed of different
substituted L-amino acids and polysaccharides. Some archaea, have cell walls composed of a
layer of proteinaceous subunits known as an S-layer, which is directly associated with the cell
membrane. This difference actually makes the Archaean members not susceptible to
antibiotics such as penicillin and lysozyme, whose antibacterial action is specifically directed to
peptidoglycan.

One of the distinctions of Archaea are their membranes, in which the lipid component of
the part contains branched isoprenes instead of fatty acids. Furthermore, these isoprenes were
joined to glycerol by ether-linkages rather than the ester linkages found in true bacteria. And
the last and the most distinctive characteristic is that Archaea are found in extreme
environments. Some extreme thermophiles can live well at over 100 C, while psychrophilic
forms of Archaea can thrive in very cold environments, like in
the Antarctic. Furthermore, other species of Archaea are
able to live in extreme alkalinity, salinity, or acidity.

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Figure 3. Triangular Disc-Shaped Halophilic Archaeon Haloarcula japonica. (Nakamura

et al. 2011)

Initially, scientists thought that archaea species were only limited to such extreme
environments. However, recent studies actually show that species of archaea can also be
observed in the world’s oceans, and also in terrestrial and semi terrestrial areas, making up a
part of the bacterial biomass. The reason that this lay undetected for so long is that these
organisms cannot as yet be cultured in the laboratory, and their presence can only be inferred
by the use of modern DNA-based analysis.

IV.2 Eukarya

The oldest known fossils are the remains of prokaryotes that lived more than 3.5 billion
years ago. Eukaryotic cells evolved more recently, about 2.5 billion years ago. According to the
endosymbiotic theory, eukaryotic cells evolved from prokaryotic cells living inside one another,
as endosymbionts (Figure 4.) In fact, the similarities between prokaryotic cells and eukaryotic
organelles provide striking evidence for this endosymbiotic relationship.

The original nucleoplasmic cell was prokaryotic. However, infoldings in its plasma
membrane may have surrounded the nuclear region to produce a true nucleus. Over time, the
chromosome of the nucleoplasm may have acquired pieces such as transposons. In some
cells, this large chromosome may have fragmented into smaller linear chromosomes. Perhaps
cells with linear chromosomes had an advantage in cell division over those with a large,
unwieldy circular chromosome. That nucleoplasmic cell provided the original host in which
endosymbiotic bacteria developed into organelles such as mitochondrion and chloroplast.
However, a major problem with this hypothesis is that it does not easily account for the fact
that Bacteria and Eukarya have similar membrane lipids, in contrast to those of Archaea. An
example of a modern prokaryote living in a eukaryotic cell is shown in Figure 5. The
cyanobacterium-like cell and the eukaryotic host require each other for survival.

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Figure 5.

Title as indicated. (Tortora et al.

2014)

Figure 4. Title as indicated. (Tortora et al. 2014).

Phylogenetic trees of species in the domain Eukarya have been constructed from
comparative sequence analysis of the 18S rRNA gene, the functional equivalent of the 16S
rRNA gene. The 18S tree shows some “early-branching” microbial eukaryotes, such as the
microsporidia and the diplomonads. By contrast, the position of these organisms on multigene
phylogenetic trees is quite different, and shows them to have arisen during a burst of
evolutionary radiation that led to most lineages of microbial eukaryotes. It is likely that this
burst in eukaryotic evolution was triggered by the onset of oxic conditions on Earth and
subsequent development of the ozone shield. The latter would have greatly expanded the
number of surface habitats available for colonization. Nevertheless, although 18S rRNA
sequencing appears to give a skewed view of eukaryotic microbial evolution, it still clearly sorts
the eukaryotes out as a distinct domain of life with evolutionary roots more closely tied to the
Archaea than to the Bacteria.

V. CLASSIFICATION AND NOMENCLATURE

Classification is the organization of organisms into progressively more inclusive groups on

the basis of either phenotypic similarity or evolutionary relationship. The hierarchical nature of

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classification is shown in Table 2. A species is made up of one to several strains, and similar
species are grouped into genera (singular, genus). Similar genera are grouped into families,
families into orders, orders into classes, up to the domain, the highest level taxon.

Table 2. Taxonomic hierarchy for the purple sulfur bacterium Allochromatium warmingii.

Nomenclature is the actual naming of organisms and follows the binomial system of
nomenclature devised by the Swedish medical doctor and botanist, Carl Linnaeus, and used
throughout biology; organisms are given genus names and species epithets. The names are
Latin or Latinized Greek derivations, often descriptive of some key property of the organism,
and are printed in italics. By classifying organisms into groups and naming them, we order the
natural microbial world and make it possible to communicate effectively about all aspects of
particular organisms, including their behavior, ecology, physiology, pathogenesis, and
evolutionary relationships. The creation of new names must follow the rules described in The
International Code of Nomenclature of Bacteria. This source presents the formal framework by
which Bacteria and Archaea are to be officially named and the procedures by which existing
names can be changed, for example, when new data warrants taxonomic rearrangements.

V. BERGEY’S MANUAL AND THE PROKARYOTES

Because taxonomy is largely a matter of scientific judgment, there is no “official”

classification of Bacteria and Archaea. Presently, the classification system most widely

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accepted by microbiologists is that of Bergey’s Manual of Systematic Bacteriology, a major

taxonomic treatment of Bacteria and Archaea. Widely used, Bergey’s Manual has served the
community of microbiologists since 1923 and is a compendium of information on all recognized
prokaryotes. Each chapter, written by experts, contains tables, figures, and other systematic
information useful for identification purposes. A second major source in bacterial diversity is
The Prokaryotes, a reference that provides detailed information on the enrichment, isolation,
and culture of Bacteria and Archaea. This work is available online by subscription through
university libraries. Collectively, Bergey’s Manual and The Prokaryotes offer microbiologists
both the concepts as well as the details of the biology of Bacteria and Archaea as we know it
today; they are the primary resources for microbiologists characterizing newly isolated
organisms.

When a new prokaryote is isolated from nature and thought to be unique, a decision
must be made as to whether it is sufficiently different from other prokaryotes to be described
as a new taxon. To achieve formal validation of taxonomic standing as a new genus or
species, a detailed description of the organism’s characteristics and distinguishing traits, along
with its proposed name, must be published, and, as just mentioned, viable cultures of the
organism must be deposited in at least two international culture collections. The manuscript
describing and naming a new taxon undergoes peer review before publication. A major vehicle
for the description of new taxa is the International Journal of Systematic and Evolutionary
Microbiology (IJSEM), the official publication of record for the taxonomy and classification of
Bacteria and Archaea. In each issue, the IJSEM publishes an approved list of newly validated
names. By providing validation of newly proposed names, publication in IJSEM paves the way
for their inclusion in Bergey’s Manual of Systematic Bacteriology.

VI. MICROBIAL SYSTEMATICS

Systematics is the study of the diversity of organisms and their relationships. It links
together phylogenyGE with taxonomy, in which organisms are characterized, named, and
placed into groups according to several defined criteria. Bacterial taxonomy traditionally has
focused on practical aspects of identification and description, activities that have relied heavily
on phenotypic comparisons. At present, the growing use of genetic information, especially
DNA sequence data, is increasingly allowing taxonomy to reflect phylogenetic relationships as
well. Bacterial taxonomy has changed substantially in the past few decades, embracing a
combination of methods for the identification of bacteria and description of new species. This
polyphasic approach to taxonomy uses three kinds of methods—phenotypic, genotypic, and
phylogenetic—for the identification and description of bacteria. Phenotypic analysis examines
the morphological, metabolic, physiological, and chemical characteristics of the cell (Table 3).
Genotypic analysis considers characteristics of the genome (Table 4). These two kinds of
analysis group organisms based on similarities. They are complemented by phylogenetic
analysis, which seeks to place organisms within an evolutionary framework.

Table 3. Some phenotypic characteristics of taxonomic value.

Category Characteristics

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Morphology Colony morphology; Gram reaction; cell size and shape; pattern of
flagellation; presence of spores, inclusion bodies (e.g., PHB,a
glycogen, or polyphosphate granules, gas vesicles, magnetosomes);
capsules, S-layers or slime layers; stalks or appendages; fruiting-body
formation
Motility Nonmotile; gliding motility; swimming (flagellar) motility; swarming;
motile by gas vesicles
Metabolism Mechanism of energy conservation (phototroph, chemoorganotroph,
chemolithotroph); utilization of individual carbon, nitrogen, or sulfur
compounds; fermentation of sugars; nitrogen fixation; growth factor
requirements
Physiology Temperature, pH, and salt ranges for growth; response to oxygen
(aerobic, facultative, anaerobic); presence of catalase or oxidase;
production of extracellular enzymes
Cell lipid chemistry Fatty acids; polar lipids; respiratory quinones
Cell wall chemistry Presence or absence of peptidoglycan; amino acid composition of
cross-links; presence or absence of cross-link interbridge
Other traits Pigments; luminescence; antibiotic sensitivity; serotype; production of
unique compounds, for example, antibiotics

Table 4. Some genotypic methods used in bacterial taxonomy.

Method Description/ Application

DNA–DNA Genome-wide comparison of sequence similarity. Useful for
hybridization distinguishing species within a genus
DNA profiling Ribotyping (Section 16.9), AFLP, rep-PCR (Figure 16.21). Rapid
method to distinguish between species and strains within a species
Multilocus sequence Strain typing using DNA sequences of multiple genes (Figure 16.22).
typing High resolution, useful for distinguishing even very closely related
strains within a species
GC ratio Percentage of guanine–cytosine base pairs in the genome. If the GC
ratio of two organisms differs by more than about 5%, they cannot be
closely related, but organisms with similar or even identical GC ratios
may be unrelated. Not much used now in taxonomy because of poor
resolution

VII. EVOLUTIONARY ANALYSIS (THEORETICAL ASPECTS)

The evolutionary history of a group of organisms is called its phylogeny, and a major goal
of evolutionary analysis is to understand phylogenetic relationships. Because we do not have
direct knowledge of the path of microbial evolution, phylogeny is inferred indirectly from
nucleotide sequence data. Our premises are that (1) all organisms are related by descent, and
(2) that the sequence of DNA in a cell’s genome is a record of the organism’s ancestry.
Because evolution is a process of inherited nucleotide sequence change, comparative

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analyses of DNA sequences allow us to reconstruct phylogenetic histories. Here, we examine

some of the ways in which this is carried out.

7.1 Genes Employed in Phylogenetic Analysis

The most widely used and useful for defining relationships in prokaryotes is the gene encoding
16S ribosomal RNA (rRNA) and its counterpart in eukaryotes, 18S rRNA, parts of the small
subunit of the ribosome.

7.2 Molecular Clocks

An unresolved question in phylogenetics is whether DNA (and protein) sequences change at a

constant rate. The approach to answering the question focuses on pairs of homologous
sequences—that is, sequences of shared evolutionary ancestry that encode functionally
equivalent molecules. If sequences do change at a constant rate, such pairs would serve as an
approximate molecular clock, allowing the time in the past when the two sequences diverged
from a common ancestral sequence to be estimated.

VIII. EVOLUTIONARY ANALYSIS (ANALYTICAL METHOD) (Madigan et al. 2013)

8.1 Obtaining DNA Sequences

Phylogenetic analysis using DNA sequences relies heavily on the polymerase chain reaction
(PCR) to obtain sufficient copies of a gene for reliable sequencing. Specific oligonucleotide
primers have been designed that bind to the ends of the gene of interest, or to DNA flanking
the gene, allowing DNA polymerase to bind to and copy the gene. The source of DNA bearing
a gene of interest typically is genomic DNA purified from particular bacterial strains, but could
be DNA extracted from an environmental sample. The PCR product is visualized by agarose
gel electrophoresis, excised from the gel, extracted and purified from the agarose, and then
sequenced, often using the same oligonucleotides as primers for the sequencing reactions.
8.2 Sequence Alignment

Phylogenetic analysis is based on homology, that is, analysis of DNA sequences that are
related by common ancestry. Once the DNA sequence of a gene is obtained, the next step in
phylogenetic analysis is to align that sequence with homologous sequences from other
organisms. By doing this, nucleotide mismatches and insertions and deletions, some of which
may be phylogenetically informative, can be pinpointed.

8.3 Phylogenetic Trees

Reconstructing evolutionary history from observed nucleotide sequence differences includes
construction of a phylogenetic tree, which is a graphic depiction of the relationships among
sequences of the organisms under study, much like a family tree. A phylogenetic tree is
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composed of nodes and branches (Figure 4). The tips of the branches represent species that
exist now and from which the sequence data were obtained. The nodes are points in evolution
where an ancestor diverged into two new organisms, each of which then began to evolve
along its separate pathway. The branches define both the order of descent and the ancestry of
the nodes, whereas the branch length represents the number of changes that have occurred
along that branch.

IX. SPECIES CONCEPT IN MICROBIOLOGY

At present, there is no universally accepted concept of species for prokaryotes. Microbial

systematics combines phenotypic, genotypic, and sequence-based phylogenetic data within a
framework of standards and guidelines for describing and identifying prokaryotes, but the issue
of what actually constitutes a prokaryotic species remains controversial.

A prokaryotic species is defined operationally as a group of strains sharing a high degree of

similarity in several independent traits. Traits currently considered most important for grouping
strains together as a species include 70% or greater genomic DNA–DNA hybridization and
97% or greater identity (3% difference) in 16S rRNA gene sequence. Experimental data
suggest that these two criteria are valid, reliable, and consistent in identifying new species of
prokaryotes. Based on genotypic criteria such as these, over 7000 species of Bacteria and
Archaea have been formally recognized.

What criteria should be used to define a genus, the next highest taxon, is more a matter of
judgment, but 16S rRNA gene sequence differences of more than 5% from all other organisms
is considered good evidence that an organism constitutes its own genus. Above the level of
genus to the family, order, and other ranks of higher taxa, no consensus ribosomal RNA
sequence-based criteria exist for delineating these ranks. Table 2 gives an example of species
definition in practice for the classification of the phototrophic purple bacterium Allochromatium
warmingii from the domain down to the species level.

How do new prokaryotic species arise? A likely possibility is by the process of periodic
purges and selection within cell populations. Imagine a population of bacteria that originated
from a single cell and that occupies a particular niche in a habitat. In theory, these cells are
genetically identical. If cells in this population share a particular resource (for example, a key
nutrient), the population is considered an ecotype (Figure 6). Different ecotypes can coexist in
a habitat, but each is only most successful within its prime niche in the habitat. However, within
each ecotype, genes mutate at random over time as the cells grow. Most of these mutations
are neutral and have no effect. However, if there is a beneficial mutation (one that increases
fitness) in a cell in one of the ecotypes, that cell will produce more progeny over time, and this
will purge the population of the original, less well-adapted cells. Repeated rounds of mutation
and selection in this ecotype lead it to become more and more distinct genetically from the
other ecotypes. Then, given enough time, cells in this lineage will carry a sufficiently large set
of unique traits that they emerge as their own species. Selection of strains bearing beneficial
mutations can proceed gradually, or it can occur quite suddenly due to rapid environmental
change. Note that this series of events within an ecotype has no effect on other ecotypes,
because different ecotypes do not compete for the same resources.

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Figure 6. A model for bacterial speciation.

X. REFERENCES

Madigan MT, Clark DP, Stahl D, Martinko JM. 2013. Brock biology of microorganisms, 13th
edition. NY, USA: Benjamin Cummings. 

Tortora G, Funke B, Case C. 2014. Microbiology: An Introduction, 11 th ed. Singapore : Pearson

Education South Asia.

https://www.researchgate.net/publication/
320413728_Difference_Between_Taxonomy_and_Systematics

https://evolution.berkeley.edu/evolibrary/article/evo_10

Imhoff JF. 2015. Thiospirillum. Bergey’s Manual of Systematics of Archaea and Bacteria, 1–
3. doi:10.1002/9781118960608.gbm01126. 

Nakamura S, Nakasone K, Takashina T. 2011. Genetics and Genomics of Triangular Disc-

Shaped Halophilic Archaeon Haloarcula japonica Strain TR-1. Extremophiles Handbook,
363–381. doi:10.1007/978-4-431-53898-1_17. 2011. 

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