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What Characteristics Have Enabled Prokaryotes to Thrive in Diverse Environments?
Small size and rapid reproduction
High genetic diversity through mutations (mutations dont occur as fast in euk, but short life span enable high genetic diversity)
Rapid evolution of diverse adaptations
Conditions on early earth made the origin of life possible
Earth formed 4.6 billion years ago, conditions were too volatile to support life
first chemical signatures of life date back to 3.6 bya
the earliest fossil evidence of organisms date to 3.5 bya (prokaryotes)
prok were alone on earth until the first eukaryote cells appeared about 1.8 bya
Chemical and physical processes on early Earth could have produced simple cells in four stages
1. Abiotic synthesis of small organic molecules
2. Joining of small molecules into macromolecules
3. Packaging of molecules into protocells, droplets with membranes that maintain consistent internal chemistry
4. Origin of self-replicating molecules
Synthesis of organoc compounds on early earth
Earths early atmosphere had little oxygen and likely contained water vapor and compounds released by volcanic erruptions
As earth cooled, water vapor condensed into oceans and most hydrogen escaped to space
Some evidence suggests that the early atmosphere was neither reducing nor oxidizing
Different models of early earth
In the 1920s, A. I. Oparin and J. B. S. Haldane hypothesized that the early atmosphere was a reducing environment
In 1953, Stanley Miller and Harold Urey showed experimentally that abiotic synthesis of organic moleculeswas possible in a reducing atmosphere
Abiotic synthesis of macromolecules
small organic molecules (monomers0 can be produced spontaneously through abiotic synthesis
amino acids and RNA nucleotides form macromolecules (polymers) without enzymes or ribosomes, when they are concentrated on hot sand, clay or rock
such polymers may have functioned as weak catalysts for chemical reactions on early earth
protocells
replication and metabolism are key properties of life that likely appeared together in early protocells
Protocells may have been fluid-filled vesicles enclosed with a membrane-like structure
In water, lipids and other organic molecules can spontaneously form vesicles with a lipid bilayer
Montmorillonite clay increases the rate of vesicle formation by concentrating molecules on its surface.
Abiotically produced vesicles can “reproduce”, “grow”, and absorb molecules from their surroundings.
Some vesicles have a selectively permeable bilayer and can perform metabolic reactions using external reagents.
Self-replicating RNA
The first genetic material was likely RNA not DNA
single stranded RNA molecules have three dimensional shapes, making catalytic function possible
Ribozymes are RNA molecules that cna catalyze reactions, some are also self replicating
Ribozyme copying errors could produce new faster repliacting shapes, those with more accurate self replication would leave more descendant molecules
In 2013, researchers built a vesicle in which RNA self replication could occur, vesciles that could grow, divide, and pass on genetic material could produce protocells that evolvd by natural selection
RNA to DNA
Life today was likely preceded by a RNA world
Single-stranded RNA could have provided the template for DNA synthesis in protocells
Double-stranded DNA is a more stable and accuratley replicated genetic material than RNA
Fossil evidence of early life
Many of the oldest fossils are stromatolites, layered rocks that formed from theactivities of prokaryotes (earliest 3.5 bya)
Stromatolites living today are frmed by cyanobacteria and other photosynthetic bacteria
Stromatolites with distinct morphologies and habitats evolved 3.1 and 2.8 bya
Stromatolites living today are formed by cyanobacteria and other photosyntheic bacteria
The cyanobacteria were the main photosyntheic organisms for over a billion years
Cyanobacteria transformed the atmosphere on early earth by reeasig oxygen produced by the photosynthesis
surving prokaryote lineages either avodied or adapted to the newly aerobic environment
Diverse structural and metabolic adaptations have eviolved in prokayotes
Most prokayotes are unicellular, some species form colonies
Prokayotic cells typically have much smaller diameters than eukaryotic cells
Prokayotes have a variety of cell shapes, including spheres (cocci), rods (bacilli), and spirals

Cell-surface structures
Cells walls protect and maintain cell shape, prevent bursting in hypotonic environments, and cause plasmolysis in hypertonic environments
Euk cell walls contain cellulose or chitin
Bacterial cell walls contain peptidoglycan, a network of modififed sugars and polypeptides
Archaeal cell wells contain polysaccarides and proteins but lack peptidogylcan
Gram positive bacteria
Bacteria have simpler cell walls compsed of a thick layer of peptidoglycan
(Biologists categorize bacteria based on cell wall composition determined by response to Gram stain)
Gram negative bacteria
Bacteria have less peptidoglycan and anouter membrane with lipopolysaccharides
Lipid portions of lipopolysaccharides can be toxic
capsule
Many antibiotics target peptidoglycan and damage bacterial cell walls
The outer-membrane protects gram-negative bacteria, increasing their antibiotic resistance
Many prokaryotes also have a sticky, protective outer layer called a capsule that allows them to adhere to the substrate or each other
Endospores
When water or nutrients are lacking, some bacteria develop resistant cells called endospores
Endospores are dormant cells that can remian viable for centuries or survive extreme conditions
Prokaryotes sticking together
Some prokaryotes stick to the substrate or each other using hairlike appendages called fimbriae
Pili (or sex pili) are longer, less numerous appendages that are used to pull prokaryotes together during DNA transfer between cells
Motility
About half of all prokaryotes exhibit taxis, the ability to move toward or away from a stimulus
EX: chemotaxis- movement from or to a chemical stimulus
Some species can move up to 50 times their body length per second
Flagella the most common motility structure in prok, likeley eveolved independently in the 3 domains
prok and euk flagella differ in structure, molecular composition, and mechanical function
The flagella of bacteria and archaea are structurally similar, but composed of unrelated proteins
Evolutionary origins of bacterial flagella
composed of three main parts (motor, hook, and filament) and 42 different proteins
many of flagellas proteins are modified versions of proteins that preform other tasks in bacteria
bacterial flagella are likely the examples of exaptation
Internal organization and DNA
prokaryote cells lack the complex compartmentalization found in euk cells
some prok have specialized infoldings of the plasma membrane that preform metabolic functions
some store metabolic products in simple protein made compartments
prok have less genomic DNA than euk, the DNA is in a singular circular chromosome in the nucleiod region
some bacteria have small rings of DNA called plasmids
prok and euk have differences in DNA replication, transcription, and translation
Nutritional and metabolic adaptations: Prokaryotes can be categorized by how they obtain energy and carbon
Phototrophs obtain energy from light
Chemotrophs obtain energy from organic compoundsor inorganic chemicals like H2S
Autotrophs use inorganic carbon sources like CO2
Heterotrophs obtain carbon from organic compounds
The role of oxygen in metabolism
Prok metabolism varies with respect to O2
Obligate aerobes require O2 for celluar respiration
Obligate anaerobes are poisioned by O2 and will use fermentation or anaerobic respiration with other substances
Facultative anaerobes can use O2 if present or not
Nitrogen Metabolism
Nitrogen is essential for the production of amino acids and nucleic acids in all organisms
• Eukaryotes cannot use atmospheric nitrogen N2
• Some prokaryotes use nitrogen fixation to convert atmospheric nitrogen N2 to ammonia NH3 that can be used by both prokaryotes and eukaryotes
Metabolic cooperation
Some prok cells cooperate to make use of the otherwise unavailable resources (ex in anabaena, photosynthetic cells exchnage products with nitrogen fixing cells called heterocysts
Cooperation between different prokaryotic species occurs in surface-coating colonies called biofilms
Biofilms are common but can cause a wide range of problems for humans, including chronic infections, tooth decay, and contamination of medical devices
Three key features of their biology allow prokaryotes to reproduce rapidly in favorable environments
Small cell size
Binary fission
Short generation times
Adaptations of prokaryotes: summary
The ongoing success of prokaryotes is primarily the result of physiological and metabolic diversification
Metabolic diversification of prokaryotes was the first great wave of adaptive radiation in the history of life
Prokaryotes have considerable genetic variation. Three factors contribute to this genetic diversity:
rapid reproduction
mutation
genetic recombination
Rapid reproduction and mutation
prok reproduce by binary fission, most offspring are identicial to parent
mutations are rare, but bc generation times are so short, they accumulate quickly in a pop
high genetic diversity allows for rapid evolution
prok are not primitive, they are highly evolved
Genetic recombination
genetic recombination is the combining of DNAs from two sources which contributes to diversity
prok DNA from different individuals can be brought together by transformation, transduction, and conjugation
movement of genes among individuals from different species is called horizontal gene transfer
Transformation and transduction
Transformation: prok cells alter their genotype with DNA taken up from the external environment
Transduction: phages (viruses that infect bacteria) move prok genes from one host cell to another
Conjugation and plasmids
Conjugation is a process by which genetic material is transfeerred directly between prok cells (E. coli exhibits this)
In bacteria, DNA transfer is one way
F factor genes can also be transferred between chromosomes during conjugation to form a recombinant bacterium
The F factor
The F factor is a piece of DNA consisting of about 25 genes required for the production of pili
it may be present in the chromosome or an f plasmid
in conjugation, cells with the f plasmid (F+) donate DNA; cells without the F plasmid (F-) recieve DNA
Transfer of the f plasmid converts an F- cell to a recombinant F+ cell
R plasmids and antibiotic resistance
Antibiotic resistance genes can be carried in R plasmids
Bacteria with resistance genes will survive exposure to antibiotics that kill bacteria without resistance genes
Antibiotic treatment increases the frequency of resistant bacteria in the population by killing the sensitive bacteria
Natural selection through widespread antibiotic use has made resistant strains of bacteria increasingly common
An overview of prokaryotic diversity
Dramatic revision of the prokaryote phylogeny has resulted from molecular research started in the 1970s (molecular analysis led to split of proks into bacteria and archaea)
New advances in molecular systematics have enabled the analysis of over 1,700 entire prokaryote genomes
Metagenomics uses the polymerase chain reaction (PCR) to sequence DNA in environmental samples (reveals the huge diversity of prok)
Molecular systematics has revealed the key role of horizontal gene transfer in the evolution of prok
Bacteria
Include the majoirty of prok familiar to people
every major mode of nutrition and metabolism is represented among bacteria, a wide variety of nutritional modes can be found even within small taxonomic groups
about 16,000 known species (estimated unknown 700,000-1.4 million)
Proteobacteria
Proteobacteria is a clade of gram-negative bacteria with diverse metabolic and nutritional modes
Evidence indicates that mitochondria evolved by ensosymbiosis from a heterotrophic proteobacterium
Chlamydias
Chlamydias are gram-negative parasites that can only live within animal host cells
Spirochetes
Spirochetes are helical gram-negative heterotrophs
They move in a spiral pattern by rotating internal, flagellum like filaments
Many species are free-living but some are parasitic
Cyanobacteria
Cyanobacteria are gram-negative photoautotrophs that generate O2 through plant-like photosynthesis
Plant chloroplasts likely evolved from cyanobacteria by the process of endosymbiosis
Cyanobacteria are abundant in freshwater and marina phytoplankton
Gram-positive bacteria
Diverse group
Actinomycetes are colony-forming bacteria composed mostly of soil decomposers and some pathogens
Streptomyces are soil dwelling bacteria that are cultured as a source of antibiotics
Other subgroups include pathogens such as Bacillus anthracis, which causes anthrax
Archaea
Archaea have many unique traits, but they also share some traits in common woith bacteria and others with eukaryotes
Some archaea are extremophiles that live in extreme environments
Extreme halophiles either tolerate or require a highly saline environment
Extreme thermophiles have adaptations that styablize DNA and proteins allowing them to thrive in extreme heat