27 - Bacteria and Archaea

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    27 - Bacteria and Archaea

    Prokaryotic Structural and Functional AdaptationsBasic Characteristics of Prokaryotes

    Most are unicellular (some remain attached after cell division)

    Cells typically .5-5 micrometers in diameter (much smaller than eukaryotes) Well organized - all life's functions in one cell

    Cell-Surface Structures

    Cell wall - protects cell, maintains shape, prevents bursting in hypotonic environments(hypertonic causes plazmolyzation, inhibiting reproduction)

    Bacterial cell walls contain peptidoglycan (polymer composed of modified sugars cross-linked by short polypeptides)

    "molecular fabric" encloses entire bacterium and anchors other molecules thatextend from its surface

    Archaeal cell walls contain various polysaccharides and proteins, but no peptidoglycan Gram stain - classifies bacterial species into two groups based on cell wall composition

    Process: Sample stained with crystal violet dye

    Sample rinsed with alcohol

    Sample stained with red dye (like safranin)

    Gram positive - simpler walls with lots of peptidoglycan, which traps crystal violetin the cytoplasm - masks red safranin, appears purple

    Gram negative - complex walls with lipopolysaccharides and very littlepeptidoglycan, so crystal violet is easily rinsed - red safranin shows, appears

    pink/red

    Medical applications: Gram negative bacteria - lipopolysaccharides are toxic (cause

    fever/shock), outer-membrane protects from body's defenses, tend to be

    more resistant to antibiotics than Gram-positive bacteria Gram positive bacteria - some virulent strains are multi-drug resistant Effectiveness of some antibiotics (penicillin) derives from their ability to

    inhibit peptidoglycan cross-linking - decreases cell wall functionality Does not harm most human cells (which have no peptidoglycan)

    Prokaryote cell wall surrounded by a capsule - sticky layer of polysaccharide or proteinwhich is dense and well defined

    Called a slime layer if it is less well organized

    Enable adherance to their substrate or other individuals in a colony

    Can protect against dehydration and the host's immune system

    Fimbriae - hairlike appendages that allow prokaryotes to stick to substrate or each other(shorter and more numerous than pili)

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    Pili (sometimes called sex pili) - appendages that pull two cells together prior to DNAtransfer from one cell to the other

    Motility

    Taxis - directed movement toward or away from a stimulus (1/2 of all prokaryotes arecapable)

    Chemotaxis - changes movement pattern in response to chemicals

    Flagella - most common motile element, either scattered over the body or concentratedat one end

    1/10 width of eukaryotic flagella; not covered by an extension of membrane

    Flagella of prokaryotes are different in molecular composition and mechanism ofpropulsion than eukaryotic flagella

    Bacterial and archaeal flagella are similar in size and mechanism, but arecomposed of different proteins

    Flagella of eukaryotes, bacteria, and archaea are most likely analogous Bacterial flagella probably evolved through exaption (was a secretory system,

    and other proteins were added over time that made it a flagellum)

    Internal Organization - DNA

    Prokaryotes lack the complex compartmentalization of eukaryotics

    Some prokaryotic cells do have specialized membranes that perform metabolic functions

    Prokaryotic genome is structurally different and usually has much less DNA

    Genome is usually a circular chromosome with far fewer proteins than eukaryotes

    Nucleoid - region of the cytoplasm that appears lighter than the surrounding cytoplasm inelectron micrographs; contains the chromosomes

    May also have plasmids - smaller rings of independently replicating DNA; generally carryonly a few genes

    Prokaryotic ribosomes tend to be smaller and differ in protein and RNA content

    Reproduction and Adaptation

    Binary fission - allows prokaryotes to reproduce very quickly in favorable environments

    Reproduction is limited by nutrient supply, the toxicity of their own metabolic waste,competition from other microorganisms, and consumption by other organisms

    Key features: Small

    Reproduce by binary fission

    Short generation times

    Can exist as large populations (in the millions)

    Biochemical and/or structural adaptations allow some prokaryotes to withstand harshenvironments

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    Endospores - resistant cells developed by some bacteria when they lackessential nutrients; original cell hides copy of chromosome in endospore, which

    lies dormant until conditions improve

    High generation rate allows fast evolution and adaptation

    Genetic Diversity in ProkaryotesRapid Reproduction and Mutation

    New mutations are rare (and generally occur by errors in DNA replication) but canincrease genetic diversity quickly in species with short generation times and large

    populations

    Diversity can lead to rapid evolution

    Genetic Recombination

    In eukaryotes, sexual process of meiosis and fertilization foster genetic recombination

    Prokaryotes use transformation, transduction, and conjugation Transformation - genotype (and possibly phenotype) of a prokaryotic cell one

    altered by the uptake of foreign DNA from the surroundings Many bacteria have cell-surface DNA-recognizing proteins

    Transduction - phages carry prokaryotic genes from one host cell to another Phage infects donor cell

    Phage DNA replicates; phage proteins halt host cell's protein synthesis

    and host cell's DNA may be fragmented

    New phages assemble, perhaps carrying host cell alleles in a phage

    capsid

    Phage carrying donor allele infects recipient cell; recombination betweendonor and recipient DNA occurs

    Genotype of resulting recombinant cell differs from donor and recipient

    Conjugation - DNA is transferred between two prokaryotic cells (usually of thesame species) that one temporarily joined

    In bacteria, DNA transfer is always one way In E. coli(best understood):

    Pilus of the donor cell attaches to recipient cell

    Pilus retracts, pulling the cells together

    Possible formation of "mating bridge" - DNA is transferred

    Ability to form pili and donate DNA due to the F (fertility) factor (piece of

    DNA)

    F plasmid - F factor in plasmid form; F+

    cells have F plasmid and

    donates DNA to F-cells (which lack the F factor); if F plasmid

    copy is transferred, F-cell becomes an F

    +cell

    Hfr (high frequency recombination) cell - F factor is built into

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    chromosome, so chromosomal genes may be transferred

    Hfr cell acts like F+

    cell

    When chromosomal DNA from an Hfr cell enters an F-cell,

    homologous regions of Hfr and F

    -

    chromosomes may align- DNA can be exchanged

    R plasmids - carry antibiotic-resistant genes

    Antibiotics kill many, but not all, bacteria

    Natural selection supports proliferation of bacteria with R plasmids

    Many can also form pili and enable conjugation

    Any are multi-antibiotic resistant

    Nutritional and Metabolic AdaptationsTrophic Types

    Phototrophs - obtain energy from light

    Chemotrophs - obtain energy from chemicals

    Autotrophs - need only CO2in some form as a carbon source

    Heterotrophs - require at least 1 organic nutrient (ex: glucose)

    Combinations lead to 4 major nutritional modes

    Mode Energy source Carbon source Type of organism

    Photoautotroph LightCO2, HCO3

    -, etc

    Photosynthetic prokaryotes

    Chemoautotroph Inorganic

    chemicalsCO2, HCO3

    -, etc

    Some prokaryotes

    Photoheterotroph Light Organic

    compounds

    Some aquatic and salt-loving

    prokaryotes

    Chemoheterotroph Organic

    compounds

    Organic

    compounds

    Many prokaryotes

    Oxygen and Metabolism

    Obligate aerobes - must use O2for cellular respiration Obligate anaerobes - poisoned by oxygen

    Some live exclusively by fermentation

    Anaerobic respiration - substances other than oxygen act as oxidizing agents inETCs

    Facultative anaerobes - use oxygen when present but carry out fermentation or

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    anaerobic respiration in an anaerobic environment

    Nitrogen Metabolism

    N is essential for the production of amino acids and nucleic acids

    Nitrogen fixation - bacteria convert atmospheric nitrogen to ammonia

    Some cyanobacteria and some methanogens (archaea) are very good - self-sufficient

    Important to other organisms (plants, particularly)

    Metabolic Cooperation

    Cells work together to perform metabolic functions not otherwise possible; generallytakes place in specialized cells of a filament

    Cyanobacteria Anabaena: some photosynthesize, others fix nitrogen

    Heterocysts - fix nitrogen; surrounded by thickened cell wall to protect from

    oxygen Intercellular connections transfer fixed nitrogen and carbohydrates

    Biofilms - surface coating prokaryotic colonies that often exhibit cooperation

    Cells secrete signaling molecules to recruit other cells

    Cells produce proteins and polysaccharides that let them stick to substrate andeach other

    Channels allow transport of nutrients and waste

    Molecular Systematics

    Genetic diversity of prokaryotes is immense Entire prokaryotic genomes can be obtained from environmental samples using

    metagenomics

    Horizontal gene transfer is significant in prokaryotic evolution

    Comparing the Three Domains of Life

    Characteristic Bacteria Archaea Eukarya

    Nuclear envelope +

    Membrane-enclosed

    organelles

    +

    Peptidoglycan in cell wall + -

    Membrane lipids Unbranched HCs Some branched

    HCs

    Unbranched

    HCs

    RNA polymerase One kind Several kinds Several kinds

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    Initiator amino acid for protein

    synthesis

    Formyl-methionine Methionine Methionine

    Introns in genes Very rare Present in some Present in many

    Response to some antibiotics Inhibited Not inhibited Not inhibited

    Histones associated with DNA In some +

    Circular chromosome + + -

    Growth at >100degrees

    Celsius In some -

    Archaea

    Extremophiles - can survive in extreme environments

    Extreme halophiles - live in highly saline environments; some tolerate the salt,others (Halobacterium) require it

    Extreme thermophiles - thrive in very hot environments

    Have adaptations in DNA and proteins that prevent denaturing

    Methanogens - release methane as a byproduct of their metabolism

    Strict anaerobes

    Live in extreme and moderate environments; swamps, guts of animals

    Decompose sewage in waste-treatment facilities

    Euryarchaeota - includes all methanogens, many extreme halophiles, and some extreme

    thermophiles Crenarchaeota - contains most extreme thermophiles

    Korarcheota - discovered in a 1996 sample in Yellowstone park

    Nanoarchaeota - very small; originally found clinging to a crenarchaeote

    Bacteria

    Proteobacteria - all gram-negative, and includes photoautotrophs, chemoautotrophs, andheterotrophs; includes anaerobes and aerobes; 5 recognized subgroups:

    Alpha proteobacteria - closely associated with eukaryotic hosts: endosymbiotic

    theory maintains that mitochondria evolved from aerobic alpha proteobacteria

    Beta proteobacteria - nutritionally diverse group including nitrogen-recyclingNitrosomonas

    Gamma proteobacteria - include sulfur-producing bacteria, Salmonella,

    Legionella, Vibrio cholerae, and E. coli

    Delta proteobacteria - include slime-secreting myxobacteria and offensive

    bdellovibrios

    Epsilon proteobacteria - most are pathogenic to humans and other animals;

    includes Campylobacter (blood poisoning, intestinal inflammation) and

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    Helicobacter pylori (ulcers)

    Chlamydias - can only survive within animal cells; gram-negative, lack peptidoglycan Chlamydia trachomatis - most common cause of blindness; most common STD

    in the US

    Spirochetes - helical heterotrophs that travel by rotation; most are free living but someare pathogenic parasites

    Treponema pallidum - syphilis Borrelia burgdorgeri - Lyme disease

    Cyanobacteria - photoautotrophs; only prokaryotes with plant-like oxygen-generatingphotosynthesis

    Endosymbiotic theory: chloroplasts likely evolved from cyanobacteria

    Gram-positive bacteria - rival proteobacteria in diversity Actinomycetes - most decompose organic matter in soil (some cause leprosy or

    TB) Bacillus anthracis - anthrax Clostridium botulinum - botulism Also includes Staphylococcus and Streptococcus Mycoplasmas - very small; only bacteria known to lack cell walls

    Prokaryotes and the BiosphereChemical Recycling

    Decomposers - break down dead organisms and waste products

    Chemoheterotrophic prokaryotes and fungi

    Prokaryotes can convert molecules to forms other organisms can use

    Cyanobacteria - use carbon dioxide to make sugars; produce oxygen Variety of prokaryotes fix atmospheric nitrogen into usable forms

    Prokaryotes can increase and decrease availability of resources for plant growth(nitrogen, phosphorus, potassium, etc)

    Ecological Interactions

    Symbiosis - two species live in close contact with each other

    Prokaryotes (symbionts) often form symbiotic relationships with larger organisms(hosts)

    Mutualism - both species benefit

    Commensalism - one species benefits, the other is not helped or harmed Parasitism - parasite eats cell contents, tissues, or body fluids of the host

    Parasites harm but usually do not kill the host (immediately)

    Pathogens - parasites that cause disease; many are prokaryotic

    Chemoautotrophic bacteria harvest energy from thermal vents to support communitieson the ocean floor, where sunlight does not reach

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    Prokaryotes and HumansMutualistic Bacteria

    Many kinds live in intestines, helping to digest various foods

    Bacterium digests food and synthesizes carbohydrates, vitamins, and other nutrients

    Signals from bacterium activate human genes which build the network of intestinal blood

    vessels necessary to absorb nutrients

    Other signals induce production of antimicrobial compounds to kill competitors

    Pathogenic Bacteria

    All known pathogenic prokaryotes are bacteria

    Some carried by pests (ex: ticks and Lyme disease

    Usually cause illness by producing poisons

    Exotoxins - proteins secreted by certain bacteria and other organisms

    Vibrio cholerae secretes exotoxin causing cholera - stimulates intestinalcells to release chloride ions; water follows by osmosis, leading to violent

    diarrhea and dehydration

    Clostridium botulinum - secretes exotoxin causing botulism as it fermentsin food

    Endotoxins - lipopolysaccharide components of the outer membrane of gram-negative bacteria (ex: Salmonella)

    Improved sanitation and antibiotics have reduced threat, but antibiotics may also helpcreate super-bacteria

    Horizontal gene transfer can make normally harmless bacteria into pathogens O157:H7 - very dangerous strain of E. coli, probably introduced by transduction

    Prokaryotes in Research and Technology

    E. coli used to clone genes

    Agrobacterium tumefaciens - helps produce transgenic crops

    PHA (polyhydroxylalkanoate) harvested from bacteria can be used in bioplastics

    Bioremidiation - use of organisms to remove pollutants from soil, air, or water

    Treating sewage, cleaning oil spills, precipitating radioactive material out ofgroundwater