Bacterial Genetics and Genetic Changeability
Bacterial Genetics
- Bacteria exhibit unique changeability compared to eukaryotes.
Barbara McClintock and Transposons
- Barbara McClintock (1902–1992) observed unpredictable kernel color inheritance in corn.
- She determined that DNA pieces moved in and out of color-related genes.
- This idea was initially met with skepticism due to the belief in DNA stability.
- By the 1970s, her idea gained acceptance, and she was awarded the Nobel Prize in 1983.
- The existence of transposons, or "jumping genes," explains the maze-like corn kernel patterns.
Staphylococcus aureus and Antibiotic Resistance
- Staphylococcus aureus is a common cause of skin and wound infections.
- Since the 1970s, it has been treated with penicillin-like antibiotics like methicillin.
- In 2004, over 60% of S. aureus strains from hospitalized patients were methicillin-resistant (MRSA).
- Millions of healthy people in the U.S. harbor MRSA.
- Healthcare-associated MRSA (HA-MRSA) is resistant to other antibiotics, including drugs of "last resort."
- Antibiotic resistance in bacteria is a significant problem due to the ability of bacteria to pass resistance genes horizontally.
Adaptation and Genetic Studies
- Organisms adapt to changing environments, with natural selection favoring those with greater fitness.
- Bacteria are an excellent system for genetic studies due to their rapid growth and large numbers.
- More is known about E. coli genetics than any other organism.
- Even a couple of cells can grow into millions of cells quickly.
- E. coli was the first organism to have its entire DNA sequenced.
Genetic Change in Bacteria
- Genetic change in bacteria occurs through mutation and horizontal gene transfer.
- Mutation involves spontaneous changes in the genome.
- Horizontal gene transfer involves the exchange of genetic information between members of the same population, such as plasmid transfer.
- Vertical gene transfer is parent to offspring.
Mutations and Phenotype
- Mutations can change an organism’s phenotype.
- Some mutants require something that other bacteria don’t need.
- If a growth factor is now required, the mutant is termed an auxotroph.
- Auxo = “increase”, troph = “nourishment”
- A prototroph does not require growth factors.
- Proto = “first”
- Geneticists compare mutants to wild type.
- New strains are designated by three-letter abbreviations, such as Trp– (cannot make tryptophan) or StrR (streptomycin resistance).
- If a bacteria can't make something anymore, it's lost the ability to make its original product.
Spontaneous Mutations
- Normal processes yield spontaneous mutations that occur randomly but infrequently.
- Mutations are passed to progeny.
- Large populations contain mutants (e.g., cells in a colony).
- A colony contains millions/billions of identical bacteria where couple mutants are similar to other bacteria.
Base Substitution
- Base substitution is the most common mutation.
- It involves the incorporation of an incorrect nucleotide during DNA synthesis.
- A point mutation is a change of a single base pair.
- This can happen because of a mistake during replication causing a complimentary change in DNA which is now stable.
Outcomes of Base Substitution
- Silent mutation: wild-type amino acid
- Missense mutation: different amino acid; resulting protein may only partially function.
- Nonsense mutation: Specifies stop codon; yields shorter protein
- Any mutation that inactivates a gene is called a knockout mutation.
Genetic Code
- A codon consists of three nucleotides read in sequence.
- Many amino acids are specified by more than one codon.
Deletion or Addition of Nucleotides
- Impact depends on number of nucleotides.
- One or two pairs yields frameshift mutation (different set of codons translated).
- Often results in premature stop codon.
- Three pairs is the same size as one codon, so it will not cause a frameshift.
- Impact depends on location within protein.
- A reading shift completely changes codons, unlike a base substitution.
Transposons (Jumping Genes)
- Can move from one location to another.
- Gene insertionally inactivated – function destroyed.
- Terminator was dance red corn-pigment gene is working.
- In others a transposon gene jumps in so it doesn't work anymore.
Transposons and Color Variation in Corn Kernels
- Classic studies carried out by Barbara McClintock observed color variation in corn kernels resulting from transposons moving into and out of genes controlling pigment synthesis.
Induced Mutations
- Result from outside influence.
- Agent that induces change is mutagen.
- Geneticists may use mutagens to increase mutation rate.
- Two general types: chemical, radiation
- Chemical or physical agents encourage mutations by 100X, 1000X.
Chemical Mutagens
- Some chemicals modify bases.
- Alkylating agents add alkyl groups onto nucleobases.
- This disrupts a bond by making a base look like another base.
- Strands of DNA look at for something side it of can.
Base Analogs
- Base analogs resemble bases.
- Can be mistakenly incorporated by DNA polymerase.
- Give bacteria something that looks like base analog.
Intercalating Agents
- Intercalating agents cause frameshift mutations.
- Flat molecules that intercalate (insert) between adjacent base pairs in DNA strand.
- Pushes nucleotides apart, produces space.
- EB showed itself between nucleotides, a mutagen that could cause error in replication.
Radiation
- Ultraviolet irradiation forms thymine dimers (covalent bonds between adjacent thymines).
- Cannot fit into double helix; distorts molecule
- Replication and transcription stall at distortion.
- Cell will die if damage not repaired.
- Mutations result from cell’s SOS repair mechanism.
- X-rays cause single- and double-strand breaks in DNA.
- Double-strand breaks often produce lethal deletions.
- Bacteria can break those T=T bonds with enzymes.
- Eukaryotes can only go in & cut out damage and fix it.
- Exposing DNA to Thymine binders: Can't repair mistakes.
- Errors = cancer.
- Break sugar phosphate backbone holding DNA when trying to repair that encourage mutations.
DNA Repair
- There is an enormous amount of spontaneous and mutagen-induced damage to DNA.
- We have mechanisms of repair so we can survive.
- If not repaired, can lead to cell death; cancer in animals.
- Spontaneous or induced mutations may be repaired by enzymes removing this.
Proofreading
- During replication, DNA polymerase sometimes incorporates wrong nucleotide.
- Mispairing slightly distorts DNA helix
- Recognized by enzymes
- Mutation prevented by repairing before DNA replication
- Two mechanisms: proofreading, mismatch repair
- Performed by DNA polymerase, verifies accuracy and can back up, excise nucleotide and incorporates correct nucleotide.
- Very efficient but not flawless
Mismatch Repair
- Fixes errors missed by DNA polymerase.
- Enzyme cuts sugar-phosphate backbone
- Another enzyme degrades short region of DNA strand
- DNA polymerase, DNA ligase make repairs
Repair of Damage from UV Light
- Photoreactivation: light repair (only found in bacteria).
- Enzyme uses energy from light to break covalent bonds of thymine dimer.
- Excision repair: dark repair (bacteria and eukaryotes).
- Enzyme removes damage; DNA polymerase and DNA ligase repair.
Mutant Selection
- Mutants are rare and difficult to isolate.
- Two main approaches:
- Direct selection: cells inoculated onto medium that supports growth of mutant but not parent (e.g., antibiotic-resistant mutants exposed to antibiotic).
- Indirect selection: isolates auxotroph from prototrophic parent strain. More difficult since parents will grow on any media on which auxotroph can grow. Replica plating allows.
Horizontal Gene Transfer
- Microorganisms commonly acquire genes from other cells through horizontal gene transfer.
- Genes naturally transferred by three mechanisms:
- Transformation: naked DNA uptake by bacteria
- Transduction: bacterial DNA transfer by viruses
- Conjugation: DNA transfer between bacterial cells
DNA Replication and Integration
- DNA replicated only if is a replicon
- Has origin of replication
- Plasmids, chromosomes
- DNA fragments added to chromosome via homologous recombination
- Only if sequence similar to region of recipient’s genome
- Cells release DNA when they are lysed
- Addition of DNase prevents transformation
Transduction
- Transfer of genes by bacteriophages
- Specialized transduction: specific genes
- Generalized transduction: any genes of donor cell
- Rare error during phage assembly
- Transfer of DNA to new bacterial host
Conjugation
- DNA transfer between bacterial cells requiring contact between donor and recipient cells
- Conjugative plasmids direct their own transfer
- F plasmid (fertility) of E. coli most studied
F Plasmids
- Encode proteins including F pilus (sex pilus) that brings cells into contact
- Enzyme cuts plasmid transferred single strand
- Complementary strands synthesized
- Both cells are now F+
Hfr Cells
- High frequency of recombination cells arise when F factor integrates into chromosome
- When an Hfr donor passes a portion of its chromosome into an F– recipient, a recombinant F– cell results.
The Mobile Gene Pool
- Genomics reveals surprising variation in gene pool of even a single species
- Perhaps 75% of E. coli genes found in all strains (core genome of species)
- Remaining make up mobile gene pool (Plasmids, transposons, genomic islands, phage DNA)
Plasmids
- Found in most Bacteria, Archaea
- Usually dsDNA with origin of replication
- Generally nonessential; cells can be cured
- Can contain few to thousands of genes
- Low or high-copy-number
- Most have narrow host range (single species); some broad host range (includes Gram– and Gram+)
Resistance Plasmids (R Plasmids)
- Resistance to antimicrobial medications, heavy metals (mercury, arsenic) compounds found in hospital environments
- Often broad host range that normal microbiota can transfer to pathogens
Transposons
- Provide mechanism for moving DNA
- Composite transposons include one or more genes
- Integrate via non-homologous recombination
VRSA
- Transposons yielded vancomycin resistant Staphylococcus aureus strain
- Patient infected with S. aureus susceptible to vancomycin
- Also had vancomycin resistant strain of Enterococcus faecalis
- Transferred transposon-containing plasmid to S. aureus
- Transposon jumped to plasmid in S. aureus