DNA Structure and Replication
DNA Structure
Definition of DNA: DNA stands for deoxyribonucleic acid.
Nature of DNA: This chemical substance is found in the nucleus of all cells in all living organisms.
Function: DNA controls all the chemical changes that occur within cells, influencing:
The type of cell that is formed (e.g., muscle, blood, nerve cells).
The kind of organism that is produced (e.g., buttercup, giraffe, herring, human).
DNA Overview
Composition of DNA:
DNA is a large biomolecule made up of long chains of subunits called nucleotides.
Each nucleotide is comprised of:
A sugar called deoxyribose.
A phosphate group denoted as -PO4.
An organic base.
Deoxyribose and Ribose
Comparison: Ribose is a sugar similar to glucose, with five carbon atoms in its molecule.
Deoxyribose: Deoxyribose resembles ribose but lacks one oxygen atom compared to ribose.
Nitrogenous Bases
Types of Nitrogenous Bases:
Purines: Contain two carbon-nitrogen rings with amino functional groups.
Pyrimidines: Contain a single carbon-nitrogen ring with amino functional groups.
Major Events in DNA Research
1869: Johann Miescher discovers an acidic substance in the nucleus containing nitrogen and phosphorus (DNA).
1928: Frederick Griffith identifies a “transforming factor” in bacteria.
1944: Avery determines that DNA is the transforming factor, though it is not widely accepted initially.
1952:
Hershey-Chase Experiment: Demonstrates that DNA is the hereditary material and not protein (widely accepted).
1952: Chargaff conducts experiments establishing the base pairings of DNA, revealing that A & T were more common than C & G and that the amounts of A/T and C/G are equal.
1952: Rosalind Franklin utilizes X-ray diffraction to obtain the first clear picture of DNA.
1953: James Watson and Francis Crick develop the molecular model of DNA, which is the double helix.
Structure of DNA
Description: DNA structure resembles a twisted ladder, known as a double helix.
Backbone Structure: Made up of deoxyribose and phosphate groups of each nucleotide.
Steps of the Ladder: Composed of nitrogenous bases (A, G, C, T).
Base Pairing: Any sequence of bases is possible, held together by hydrogen bonds.
Historical Context: Other models of DNA were proposed, but only the double helix model met both Chargaff’s data and Franklin’s X-ray observations.
Incorrect DNA Structure Models
Triple Helix Model: Proposed by Linus Pauling just prior to Watson and Crick's model.
Flaws: Incorporates phosphate groups on the inside, with nucleotides pointing outward, which results in:
Negatively charged phosphates repelling each other.
Weak interactions that do not adequately hold the structure together.
Directionality of DNA
Strand Directionality:
Phosphate on the 5’ carbon of one nucleotide bonds to the 3’ carbon of the next nucleotide.
This establishes a direction for each DNA strand, starting from 5’ end to 3’ end.
The strands in the double helix are antiparallel:
One strand runs 5’ to 3’ while the complementary strand runs 3’ to 5’.
Anti-parallel Strands of DNA
Bonding in the DNA Backbone:
Nucleotides are covalently bonded via phosphodiester bonds between the 3’ and 5’ carbons, giving the DNA molecule a defined direction.
Each complementary strand runs in the opposite direction:
One strand: 5’ to 3’.
Complementary strand: 3’ to 5’.
Chargaff’s Rule
Chargaff's Analysis: Analyzed DNA across various organisms and found:
Adenine (A) always equals Thymine (T).
Guanine (G) always equals Cytosine (C).
Significance: Establishes that a purine always bonds to a pyrimidine.
Chromosome Structure in Prokaryotes
Characteristics:
Example: E. coli bacterium.
Structure: The DNA molecule is a single, double-stranded circular loop.
Replication process is generally similar across different DNA structures.
Size: Contains approximately 5 million base pairs and 3,000 genes.
DNA Replication: General Overview
Purpose: Cells must replicate DNA before division to ensure each daughter cell inherits a complete genetic copy.
Models of Replication:
Semiconservative: Each strand serves as a template.
Conservative: Entire DNA molecule acts as a template.
Dispersive: Existing strands are broken and mixed with new segments.
Meselson and Stahl Experiment
Experiment Outline: Bacteria cultured in a medium containing heavy nitrogen (15N) were transferred to a medium with lighter nitrogen (14N).
Observations:
After the first replication, a mixture of heavy and light DNA was seen (indicative of semi-conservative replication).
Following the second replication, lighter DNA predominated.
Semi-Conservative Model
Mechanism: During DNA replication,
Each strand serves as a template, resulting in new strands comprised of half parental template and half new DNA.
Steps in DNA Replication
Coordination by Enzymes: A variety of enzymes work together at the replication fork.
Similar Processes: Occurs in both eukaryotes and prokaryotes.
Initial Unwinding:
DNA Gyrase: Unwinds supercoiled DNA.
Helicase Enzyme: Separates hydrogen bonds between base pairs.
Single-Stranded Binding Proteins: Stabilize unwound DNA, preventing reformation of hydrogen bonds.
RNA Primer Addition:
Primase: Adds a small section of RNA known as the RNA primer at the 3' end of the template DNA.
In eukaryotes, primase operates with DNA polymerase α to synthesize DNA-RNA primers.
Building New Strands:
DNA Polymerase III: Synthesizes new daughter strands by adding complementary bases to the parent strands.
In eukaryotes, DNA polymerase δ interacts with Proliferating Cell Nuclear Antigen (PCNA), acting as a sliding clamp.
Completing the Replication:
DNA Polymerase I: Removes RNA primers and fills in the gaps with DNA in prokaryotes. Similarly completed by DNA polymerase δ in eukaryotes.
DNA Ligase: Joins fragments of DNA into a cohesive strand that is then wound tightly into structure.
Okazaki Fragments
Observation: Both DNA strands must be replicated in both directions. Since DNA polymerases can only operate in a 5’ to 3’ direction, this leads to:
Leading Strand: Replicated continuously in one long pass.
Lagging Strand: Replicated in short segments known as Okazaki fragments.
Overall Summary of Replication Steps**
Helicase: Separates the DNA strands.
Single-Stranded Binding Proteins (SSB): Prevent re-annealing.
Primase: Synthesizes RNA primers.
DNA Polymerase I: Removes RNA primers and fills in gaps.
DNA Ligase: Joins DNA fragments together.
Leading & Lagging Strands: Results in one leading strand and one lagging strand due to directionality constraints.