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