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apter 26: Bacteria and Archaea: the Prokaryotic Domains TER 26 Bacteria and Archaea: The Prokaryotic Domains Mycobacterium tuberculosis Color-enhanced images shows rod-shaped bacterium responsible for tuberculosis (Raven et al 2002) Nitrogen cycle Endosymbiotic Theory

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Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

TER 26

Bacteria and Archaea:The Prokaryotic Domains

Mycobacterium tuberculosis Color-enhanced images shows rod-shaped bacterium responsible for tuberculosis (Raven et al 2002)

Nitrogen cycle

Endosymbiotic Theory

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Structure of a Prokaryotic Cell

Structure of a Eukaryotic Animal Cell

Structure of a Eukaryotic Plant Cell

Prokaryotic cells have a simple interior organization compared to Eukaryotes.

•Membrane-enclosed nucleus lacking

•Membrane-enclosed cytoplasmic organelles lacking

•Cytoskeleton lacking-support from rigid cell wall

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Lecture Themes

•origins, evolution and diversity

•structure and function

•ecological function and relationships

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

1. Prokaryote Phylogeny

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Genome of the Archaeon Methanococcus jannaschii was sequenced in 1996. Sequencing of M. jannashcii confirmed Carl Woese’s long-standing hypothesis that life traces back to three main lineages, one of which (Archaea) includes prokaryotes that share a more recent common ancestry with eukaryotes than with the prokaryotic “true bacteria”

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Prokaryotic Structure and Function

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Cyanobacteria 10 um dia.

E. coli 1X2 um

Mycoplasma 0.3-0.8 um dia.

Bacteriophage 0.07X 0.2 um

Viroid 0.01 X 0.3 um

Lymphocycte 10 um dia.

Paramecium 30X 75 um

Sizes of viruses, bacteria and eukaryotes compared Most bacteria are 1-5 um diameter (most Eukaryotic cells are 10-100 um)

Bacillus on the head of a pin

(Keaton 1993)

Largest known prokaryote is the marine bacterium Thiomargarita namibiensis; bright white cell in upper left, about .75 mm dia., attached to two dead ones. Fruitfly in picture for size comparison.

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Pseudomonas aeruginosa

Streptococcus Spirillum volutans

Raven et al 2002

Spherical coccus (Enterococcus)

Rod-shaped bacillus(E. coli)Bacterial Form

Three shapes are especially common among bacteria – spheres, rods and spirals

Most are unicellular, some aggregate transiently, some form permanent aggregations of identical cells;some show division of labor between two or more specialized cell times

Helical spirilla(|Aquaspirillum spirosa)

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Scanning electron micrograph of a colony of streptomyces, one of the actinomycetes. The actinomycetes have a much more complicated morphology than most other bacteria. (Keaton and Gould 1993)

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

The exterior surfaces of Prokaryotes. Almost all prokaryotes have a cell wall, and in most that wall contains peptidoglycan – polymers of modified sugars that are cross-linked by short polypeptides

•Most bacterial cell walls contain peptidoglycan (lacking in Archaea)

•Gram staining is an important technique for identifying bacterial; cells stain differentially based on structure and composition of walls

•Pathogenesis is related to cell wall structure and composition

•Many antibiotics act by preventing formation of cell walls, by inhibiting synthesis of cross-links in peptidoglycan

•Many prokaryotes produce capsules that function in adherance and protection

•Many prokaryotes have surface appendages called pili that are function in adherance Penicillium

chrysogenumNeisseria gonorrhoeae

E. coli

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Mechanisms of Motility Many bacteria are motile. Fllagellar action is the most common,but not the only mechanism, for generating movement.

•Prokaryotic flagella

•Flagella-like helical filaments

•Growing gelatinous threads

Motility Behavior

•Kinesis

•Taxis

Spirillum volutans Borrelia burgdorferi

Aquaspirillum sinosum

Lyme disease symptoms, and the disease vector – a tick

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Electron micrograph of E. coli shoing long helical flagella.

1 um

0.05 um

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Bacteria swim by rotating their flagella.

Vibrio cholerae (pathogen responsbible for cholera); the unsheathed core visible at top of photo is composed of a single crystal of the protein flagellin.

In intact flagella, core is surrounded by a flexible sheath. Rotary motion of the motor creates a kind of rotary motion when organism swims.

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Exensive folded photosynthetic membranes are visisble in Prochloron cell. The single, circular DNA molecule is located in the clear area in the central region of the cell.

The mesosome is an infolding of the plasma membrane serves as a point of attachment for DNA in some bacterial cells

mesosome

plasma membrane

DNA

•various specialized membranes, but lacking extensive compartmentalization by internal membranes

•ribosomes present but differ from eukaryotic ones in size and composition

•genomes are smaller and simpler than in eukaryotes; one major chromosome and, in some species, plasmids

•Processes of DNA replicatin and protein translation are generally similar to eukaryotes

Cellular and Genomic Organization The organization of cellular components, including the genome, differs substantially between prokaryotes and eukaryotes

Infoldings of plasma membrane, similar in ways to cristae of mitochondria, function in cellular respiration in aerobic bacteria

Thylakoid membranes of photosynthetic cyanobacteria

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Prokaryote Reproduction and Population Growth Prokaryote populations grow and adapt rapidly, through asexual reproduction as well as mechanisms involving gene transfer

Cell divisionAsexual reproduction by cell division via binary fission

Mechanisms of gene transfer-transformation; genes from environment-conjugation; genes from another prokaryote-transduction genes via a virus

Adaptationshort generation time allows favorable mutations and novel genomes arising from gene transfer to spread quickly in rapidly reprducing

Growth virtual geometric growth while in environments with unlimited resources

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Dormancy and Endosporulation Some bacteria form highly resistant spores under harsh environmental conditions

Antibiotic synthesisSome prokaryotes (and protists and fungi) synthesize and release antibiotic chemicals that inhibit growth of other microbes

Adaptations to Harsh Environmental Conditions: Some bacteria are capable of dormancy, endosporulation and antibiotic synthesis

Bacillus anthracus

Sporulating Bacillus cell

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Nutritional and Metabolic Diversity

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Nutrition; how an organism obtains two resources from the environment;

-energy

-carbon source to build the organic molecules of cells

•Phototrophs; use light energy

•Chemotrophs; obtain energy from chemicals taken from the environment

•Autotroph; needs only the inorganic compound CO2 as a carbon source

Hetertroph: requires at least one organic nutrient for making other organic compounds

Sources: Campbell et al (2002), Freeman (2002), Purves et al (2001)

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

The basic themes of metabolism, among all domains are

-extracting usable energy from reduced compounds -using light to produce high-energy electrons-fixing carbon.

All organisms have mechanisms for trapping usable energy in ATP; ATP allows cells to do work; there is no life without ATP

At one point or another, you have studied these metabolic themes as they occur Eukaryotes and perhaps Prokaryotes; photosynthesis(eg,in green plants and respiration (eg in all Eukaryotes)

Prokaryotes show tremendous diversity in metabolic process.in that they have evolved dozens of variations on these most basic themes of metabolism

This Prokarotic metabolic diversity is important for two reasons:

1.It explains their ecological diversity; they are found almost everywhere because they exploit such a tremendous variety of molecules as food

2.Global nutrient cycling of (eg nitrogen, phosphorous, sulfur, carbon) is mediated by, exists because, prokaryotes can use them in almost any molecular form

Sources: Freeman 2002, Campbell 2002

Overview of photosynthesis and respiration

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Overview of cellular respiration One (very important!!) example of metabolic pathways by which many species obtain energy for generating ATP by oxidizing reduced organic compounds

Highly reduced molecule, glucose, serves as original electron donor (ie, molecule is oxidized) and highly oxidized molecule, oxygen, serves as final electron acceptor

Overview of Photosynthesis

Many prokaryotes generate ATP by employing electron donors and acceptors other than sugars and oxygen, and produce by-products other than water

Sources: Freeman 2002, Campbell 2002

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Some Electron Donors and Acceptors Used by Bacteria and Archaea

Electron Donor Electron Acceptor Product Metabolic Strategy *

H2 or organic compounds SO42- H2S sulfate-reducers

H2 CO2 CH4 methanogens

CH4 O2 CO2 methanotrophs

S or H2S O2 SO42- sulfur bacteria

organic compounds Fe3+ Fe2+ iron-reducers

NH3 O2 NO2- nitrifiers

organic compounds NO3- N2O, NO or N2 denitrifiers (or nitrate reducers)

NO2- O2 NO3

- nitrosifiers

* This column gives the name biologists use to identify species that use a particular metabolic strategy

Source: Freeman (2002), Purves et al (2001)

nitrification: oxidation of ammonia to nitrite and nitrate ions

denitrification: reduction of nitrogen-containing ions to form nitrogen gas and other products

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

•lateral gene transfer has taken place repeatedly through transformation and viral infection

•in transfers among bactera and archaea the primary mechanism probably involves loops of mobile DNA (plasmids)

•swapped genes tend to be those involved in energy and carbon metabolism (not information processing , eg DNA replication, transcription, protein synthesis) – interesting…as metabolic diversity is a hallmark of the Bacteria and Archaea!!

Source: Freeman (2002)

Lateral Gene Transfer. Gray branches show diversification of the three domains. Red branches show movement of genes from species in one part of the tree to species in other parts

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Nutritional Diversity among Chemoheterotrophs (most known Prokaryotes)

•Saprobes; decomposers that absorb nutrients from dead organic matter

•Parasites; absorb nutrients from body fluids of living hosts

Relevance of Oxygen to Metabolism among Bacteria and Archaea

•Obligate aerobes

•Facultative anaerobes

•Obligate anaerobes

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Chapter 26: Bacteria and Archaea: the Prokaryotic DomainsChapter 26: Bacteria and Archaea: the Prokaryotic Domains

Why Three Domains?Why Three Domains?

General Biology of the ProkaryotesGeneral Biology of the Prokaryotes

Prokaryotes in Their EnvironmentsProkaryotes in Their Environments

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Prokaryote Phylogeny and DiversityProkaryote Phylogeny and Diversity

The BacteriaThe Bacteria

The The ArchaeaArchaea

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Why Three Domains?

• Living organisms can be divided into Living organisms can be divided into three domains: Bacteria, Archaea, and three domains: Bacteria, Archaea, and Eukarya. The prokaryotic Archaea and Eukarya. The prokaryotic Archaea and Bacteria differ from each other more Bacteria differ from each other more radically than the Archaea from the radically than the Archaea from the Eukarya.Eukarya.

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Why Three Domains?

• Evolutionary relationships of the Evolutionary relationships of the domains were revealed by rRNA domains were revealed by rRNA sequences. Their common ancestor sequences. Their common ancestor lived more than 3 billion years ago, lived more than 3 billion years ago, that of the Archaea and Eukarya at that of the Archaea and Eukarya at least 2 billion years ago. Review Figure least 2 billion years ago. Review Figure 26.226.2 and Table and Table 26.126.1

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Figure 26.2

Figure 26.2Figure 26.2

figure 26-02.jpg

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Table 26.1

Table 26.1Table 26.1

table 26-01.jpg

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

General Biology of the Prokaryotes• The prokaryotes are the most The prokaryotes are the most

numerous organisms on numerous organisms on Earth,occupying an enormous variety Earth,occupying an enormous variety of habitats.of habitats.

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

General Biology of the Prokaryotes • Most prokaryotes are cocci, bacilli, or Most prokaryotes are cocci, bacilli, or

spiral forms. Some link together to spiral forms. Some link together to form associations, but very few are form associations, but very few are truly multicellular.truly multicellular.

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

General Biology of the Prokaryotes • Prokaryotes lack nuclei, membrane-Prokaryotes lack nuclei, membrane-

enclosed organelles, and enclosed organelles, and cytoskeletons. Their chromosomes are cytoskeletons. Their chromosomes are circular. They often contain plasmids. circular. They often contain plasmids. Some contain internal membrane Some contain internal membrane systems.systems.

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

General Biology of the Prokaryotes • Many prokaryotes move by means of Many prokaryotes move by means of

flagella, gas vesicles, or gliding flagella, gas vesicles, or gliding mechanisms. Prokaryotic flagella mechanisms. Prokaryotic flagella rotate.rotate.

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

General Biology of the Prokaryotes • Prokaryotic cell walls differ from those Prokaryotic cell walls differ from those

of eukaryotes. Bacterial cell walls of eukaryotes. Bacterial cell walls generally contain peptidoglycan. generally contain peptidoglycan. Differences in peptidoglycan content Differences in peptidoglycan content result in different reactions to the result in different reactions to the Gram stain. Review Figure Gram stain. Review Figure 26.726.7

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Figure 26.7

Figure 26.7Figure 26.7

figure 26-07.jpg

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

General Biology of the Prokaryotes • Prokaryotes reproduce asexually by Prokaryotes reproduce asexually by

fission, but also exchange genetic fission, but also exchange genetic information.information.

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

General Biology of the Prokaryotes • Prokaryotes’ metabolic pathways and Prokaryotes’ metabolic pathways and

nutritional modes include obligate and nutritional modes include obligate and facultative anaerobes, and obligate facultative anaerobes, and obligate aerobes. Nutritional types include aerobes. Nutritional types include photoautotrophs, photoheterotrophs, photoautotrophs, photoheterotrophs, chemoautotrophs, and chemoautotrophs, and chemoheterotrophs. Some base chemoheterotrophs. Some base energy metabolism on nitrogen- or energy metabolism on nitrogen- or sulfur-containing ions. Review Figure sulfur-containing ions. Review Figure 26.826.8 and Table and Table 26.226.2

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Figure 26.8

Figure 26.8Figure 26.8

figure 26-08.jpg

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Table 26.2

Table 26.2Table 26.2

table 26-02.jpg

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Prokaryotes in Their EnvironmentsProkaryotes in Their EnvironmentsSome prokaryotes play key roles in global nitrogen and sulfur cycles. Nitrogen Some prokaryotes play key roles in global nitrogen and sulfur cycles. Nitrogen fixers, nitrifiers, and denitrifiers do so in the nitrogen cycle. Review Figure fixers, nitrifiers, and denitrifiers do so in the nitrogen cycle. Review Figure 26.1026.10

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Figure 26.10

Figure 26.10Figure 26.10

figure 26-10.jpg

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Prokaryotes in Their Environments • Photosynthesis by cyanobacteria Photosynthesis by cyanobacteria

generated the oxygen gas that generated the oxygen gas that permitted the evolution of aerobic permitted the evolution of aerobic respiration and the appearance of respiration and the appearance of present-day eukaryotes.present-day eukaryotes.

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Prokaryotes in Their Environments • Many prokaryotes live in or on other Many prokaryotes live in or on other

organisms, with neutral, beneficial, or organisms, with neutral, beneficial, or harmful effects.harmful effects.

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Prokaryotes in Their Environments • A minority of bacteria are pathogens. A minority of bacteria are pathogens.

Some produce endotoxins, which are Some produce endotoxins, which are rarely fatal; others produce often rarely fatal; others produce often highly toxic exotoxins.highly toxic exotoxins.

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Prokaryote Phylogeny and Diversity• Phylogenetic classification of Phylogenetic classification of

prokaryotes is based on rRNA prokaryotes is based on rRNA sequences and other molecular sequences and other molecular evidence.evidence.

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Prokaryote Phylogeny and Diversity • Lateral gene transfer among Lateral gene transfer among

prokaryotes makes it difficult to infer prokaryotes makes it difficult to infer prokaryote phylogeny. prokaryote phylogeny.

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Prokaryote Phylogeny and Diversity • Evolution can proceed rapidly in Evolution can proceed rapidly in

prokaryotes because they are haploid prokaryotes because they are haploid and can multiply rapidly.and can multiply rapidly.

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

The Bacteria

• There are far more known bacteria There are far more known bacteria than archaea. One phylogenetic than archaea. One phylogenetic classification of the domain Bacteria classification of the domain Bacteria groups them into over a dozen groups.groups them into over a dozen groups.

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

The Bacteria

• The most ancient bacteria, like the The most ancient bacteria, like the most ancient archaea, may be most ancient archaea, may be thermophiles, suggesting that life thermophiles, suggesting that life originated in a hot environment.originated in a hot environment.

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

The Bacteria

• All four nutritional types occur in the All four nutritional types occur in the Proteobacteria. Metabolism in different Proteobacteria. Metabolism in different proteobacteria groups has evolved proteobacteria groups has evolved along different lines. Review Figure along different lines. Review Figure 26.1226.12

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Figure 26.12

Figure 26.12Figure 26.12

figure 26-12.jpg

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

The Bacteria

• Cyanobacteria, unlike other bacteria, Cyanobacteria, unlike other bacteria, photosynthesize using the same photosynthesize using the same pathways plants use. Many fix pathways plants use. Many fix nitrogen.nitrogen.

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

The Bacteria

• Spirochetes move by means of axial Spirochetes move by means of axial filaments.filaments.

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

The Bacteria

• Chlamydias are tiny parasites that live Chlamydias are tiny parasites that live within the cells of other organisms.within the cells of other organisms.

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

The Bacteria

• Firmicutes are diverse; some produce Firmicutes are diverse; some produce endospores, resting structures endospores, resting structures resistant to harsh conditions. Some resistant to harsh conditions. Some actinomycetes produce important actinomycetes produce important antibiotics. Actinomycetes grow as antibiotics. Actinomycetes grow as branching filaments.branching filaments.

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

The Bacteria

• Mycoplasmas, the tiniest living things, Mycoplasmas, the tiniest living things, lack conventional cell walls and have lack conventional cell walls and have very small genomes.very small genomes.

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

The Archaea

• Archaea cell walls lack peptidoglycan, Archaea cell walls lack peptidoglycan, and their membrane lipids contain and their membrane lipids contain branched long-chain hydrocarbons branched long-chain hydrocarbons connected to glycerol by ether connected to glycerol by ether linkages. Review Figure linkages. Review Figure 26.2226.22

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

Figure 26.22

Figure 26.22Figure 26.22

figure 26-22.jpg

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

The Archaea

• The domain Archaea can be divided The domain Archaea can be divided into two kingdoms: Crenarchaeota and into two kingdoms: Crenarchaeota and Euryarchaeota.Euryarchaeota.

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

The Archaea

• Crenarchaeota are heat-loving and Crenarchaeota are heat-loving and often acid-loving archaea.often acid-loving archaea.

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

The Archaea

• Methanogens produce methane by Methanogens produce methane by reducing carbon dioxide. Some live in reducing carbon dioxide. Some live in the guts of herbivorous animals; some the guts of herbivorous animals; some in high-temperature environments on in high-temperature environments on the ocean floor.the ocean floor.

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

The Archaea

• Extreme halophiles are salt lovers that Extreme halophiles are salt lovers that lend a pinkish color to salty lend a pinkish color to salty environments; some grow in extremely environments; some grow in extremely alkaline environments.alkaline environments.

Chapter 26: Bacteria and Archaea: the Prokaryotic Domains

The Archaea

• Archaea of the genus Thermoplasma Archaea of the genus Thermoplasma lack cell walls, are thermophilic and lack cell walls, are thermophilic and acidophilic, and have a tiny genome acidophilic, and have a tiny genome (1,100,000 base pairs).(1,100,000 base pairs).