Molecular Biology/ Table of Contents/ Chapter 4
⌂ HomeTable of contents
Chapter 4

DNA, Chromosomes, and Genomes

Molecular Biology of the Cell · Part 1 of 2 · 10 MCQs per part
Part 1 of 2
DNA Structure, the Nucleosome, and Chromatin Packaging
The two meters of DNA in a human cell must be compacted 10,000-fold to fit into a nucleus 6 μm across — yet specific genes must remain accessible to the transcription machinery at any moment. Understanding how DNA is packaged explains gene regulation, inheritance, and the origins of cancer.

4.1 The DNA Double Helix

DNA is a double helix composed of two antiparallel polynucleotide strands wound around each other. Each strand is a polymer of deoxyribonucleotides linked by 3′–5′ phosphodiester bonds. The four bases — adenine (A), thymine (T), guanine (G), and cytosine (C) — project inward and pair specifically: A with T (two hydrogen bonds) and G with C (three hydrogen bonds). This complementary base pairing means the sequence of one strand dictates the other, enabling faithful replication and transcription.

The helix has a major groove and a minor groove, which proteins exploit for sequence-specific DNA recognition. Under physiological conditions, DNA adopts the B-form helix with ~10.5 base pairs per turn and a rise of ~3.4 Å per base pair.

Key term
Complementary base pairing

The specific hydrogen-bonding between A–T and G–C pairs that holds the two strands of a DNA double helix together and governs replication, transcription, and translation fidelity.

4.2 Nucleosomes: First Level of Compaction

DNA in eukaryotes wraps around histone octamers to form nucleosomes — the fundamental repeating unit of chromatin. Each octamer contains two copies each of histones H2A, H2B, H3, and H4. About 147 bp of DNA makes ~1.65 left-handed turns around the histone core. Adjacent nucleosomes are connected by linker DNA (10–80 bp) and can be compacted further with the help of linker histone H1.

This arrangement reduces DNA length by about 7-fold. The N-terminal histone tails protrude from the nucleosome surface and are subject to extensive post-translational modifications (acetylation, methylation, phosphorylation, ubiquitylation) that regulate chromatin structure and gene expression.

Key term
Nucleosome

The basic structural unit of chromatin: 147 bp of DNA wound ~1.65 turns around an octamer of histone proteins (two each of H2A, H2B, H3, H4), reducing DNA length ~7-fold.

4.3 Higher-Order Chromatin Structure

Nucleosomal arrays compact further into a 30-nm fiber (though its exact structure remains debated), and then into larger loops, domains, and ultimately the highly condensed chromosomes visible at mitosis. Chromatin exists in two functional states: euchromatin — relatively open, gene-rich, and transcriptionally active — and heterochromatin — densely packed, gene-poor, and transcriptionally silent. Heterochromatin is further divided into constitutive heterochromatin (e.g., centromeres, telomeres) and facultative heterochromatin (e.g., the inactive X chromosome).

Pause & Recall
Why do histone modifications not directly alter DNA sequence, yet can still be inherited through cell division?
Histone modifications constitute epigenetic information. Enzymes that write modifications (e.g., histone methyltransferases) can be recruited to daughter nucleosomes by modified parental histones, propagating the modification pattern after replication. Some modifications recruit chromatin-modifying complexes that reinforce the same state, creating a self-sustaining epigenetic mark without changing the DNA sequence.
Practice Questions — Part 1Score: 0 / 10

1. In the DNA double helix, which bases pair with each other?

2. How many base pairs of DNA wrap around a single histone octamer?

3. The histone octamer in a nucleosome core particle is composed of:

4. Euchromatin differs from heterochromatin in that euchromatin is:

5. Which groove of the DNA double helix is primarily used for sequence-specific protein recognition?

6. The two strands of the DNA double helix run in what orientation relative to each other?

7. Which modification to histone H3 lysine 9 (H3K9me3) is most associated with:

8. Linker histone H1 functions primarily to:

9. The B-form DNA helix has approximately how many base pairs per turn?

10. Centromeres are an example of:

Part 1 complete! Score: 0 / 10

Section B · Recall Questions · Part 1

Type your answer, then click Check to reveal the sample answer.

B1

Describe the key structural features of the DNA double helix.

B2

What is a nucleosome and how does it contribute to DNA compaction?

B3

Name two types of histone modifications and describe how they generally affect gene expression.

B4

Distinguish between constitutive and facultative heterochromatin with one example of each.

B5

Explain why the major groove of DNA is important for protein–DNA recognition.

B6

What is DNA methylation at CpG dinucleotides and how does it relate to gene silencing?

B7

Why are topoisomerases needed during DNA replication and transcription?

B8

What are telomeres and why are they essential for chromosome integrity?

B9

What is the function of the centromere during cell division?

B10

State Chargaff's rules and explain what they imply about DNA structure.

Section C · Critical Thinking · Part 1

Analyze and apply concepts; compare your reasoning to the sample answer.

C1

A drug inhibits histone deacetylases (HDACs). Predict the effect on chromatin structure and gene expression, and explain why HDAC inhibitors are investigated as anticancer agents.

C2

How can a differentiated cell (e.g., a liver cell) maintain its identity through hundreds of cell divisions without changing its DNA sequence?

C3

Why can't transcription factors simply bind to their target sequences in nucleosomal DNA without assistance? What machinery helps them gain access?

C4

Most cancer cells reactivate telomerase. Explain why this is essential for tumor progression and why normal somatic cells suppress telomerase.

C5

What are topologically associating domains (TADs) and why is their disruption clinically significant?

Section D · Interactive Questions · Part 1

Enter your answer and click Check for instant feedback.

D1

How many hydrogen bonds hold an A–T base pair together? (number)

D2

How many base pairs of DNA are wrapped around each histone octamer? (number)

D3

What histone variant replaces H3 at centromeres? (one word, all caps)

D4

The human telomere repeat sequence is TTAGGG. How many guanines are in each repeat? (number)

D5

What enzyme adds telomeric repeats to chromosome ends? (one word)
Part 2 →
You've covered DNA structure and chromatin packaging. Part 2 explores how genomes are organized — the types of sequences they contain, how gene families arise, and what transposable elements reveal about genome evolution.
Part 2 of 2
Genome Organization, Gene Families, and Transposable Elements
Human DNA contains ~3.2 billion base pairs, but only ~1.5% encodes proteins. The rest — once dismissed as "junk" — turns out to include regulatory elements, structural sequences, and the remnants of millions of transposable element insertions that shaped the genome over evolutionary time.

4.4 Genome Organization and Repetitive DNA

Eukaryotic genomes contain a mix of unique sequences (single-copy genes), moderately repetitive sequences (gene families, tandem repeats such as rRNA and histone genes), and highly repetitive sequences (satellite DNA at centromeres/telomeres, which reassociates very rapidly in renaturation experiments — Cot analysis). In humans, about 50% of the genome derives from transposable elements.

C-value paradox: genome size (C-value) does not correlate with organismal complexity. Onions have ~5× more DNA than humans. This reflects differences in the amount of non-coding and repetitive DNA, not gene number.

Key term
C-value paradox

The lack of correlation between genome size (C-value, the haploid DNA content) and organismal complexity, explained by variable amounts of non-coding repetitive sequences across species.

4.5 Gene Families and Genome Duplication

Many genes exist in related copies called gene families that arose by gene duplication and divergence. Examples include the globin family (α- and β-globin clusters), Hox genes, and immunoglobulin genes. After duplication, one copy can evolve new functions (neofunctionalization) or the two copies can subdivide ancestral functions (subfunctionalization). Some duplicates become pseudogenes — nonfunctional remnants bearing mutations that disable expression.

4.6 Transposable Elements

Transposable elements (TEs) are DNA sequences that can move within the genome. In humans, most TEs are non-autonomous remnants. They are classified as:

Class I (retrotransposons): move via an RNA intermediate ("copy-and-paste"). LINEs (Long Interspersed Elements, especially LINE-1) and SINEs (Short Interspersed Elements, especially Alu) are the most abundant human TEs.

Class II (DNA transposons): move via "cut-and-paste" using transposase, without an RNA intermediate. These are largely inactive in humans.

TEs have shaped genomes by creating mutations, new regulatory elements, novel exons (exon shuffling), and species-specific gene expression patterns. Active retrotransposition can cause disease (e.g., LINE-1 insertions in cancer).

Pause & Recall
What is the difference between a gene family and a pseudogene?
A gene family consists of related, functional genes that arose by duplication and divergence, each producing a protein with similar but not identical function (e.g., α- and β-globins). A pseudogene is a nonfunctional copy of a gene that has accumulated mutations (stop codons, frameshifts, lack of regulatory sequences) that prevent it from being expressed or producing functional protein. Pseudogenes represent evolutionary dead ends, while gene family members are retained by selection.
Practice Questions — Part 2Score: 0 / 10

1. The C-value paradox refers to the observation that:

2. LINEs and SINEs are classified as:

3. A pseudogene differs from a functional gene family member in that it:

4. Gene duplication followed by mutation of one copy to acquire a new function is termed:

5. What fraction of the human genome consists of protein-coding sequences?

6. The globin gene family in humans is an example of:

7. Highly repetitive satellite DNA is characterized by:

8. Transposable elements have contributed to genome evolution by:

9. The most abundant short repetitive element in the human genome (~11% of human DNA) is:

10. Which best describes a processed pseudogene?

Part 2 complete! Score: 0 / 10

Section B · Recall Questions · Part 2

Type your answer, then click Check to reveal the sample answer.

B1

Describe how a retrotransposon moves within the genome.

B2

How do multigene families arise and what evolutionary advantage do they provide?

B3

What are Hox genes and why is their chromosomal organization significant?

B4

Why are rRNA genes present in hundreds of copies in most eukaryotic genomes?

B5

What are long non-coding RNAs (lncRNAs) and give one example of their functional role.

B6

How can unequal crossing over between tandem repeats lead to copy-number variation and disease?

B7

What is genomic imprinting and why does it violate Mendelian expectations?

B8

What is a single-nucleotide polymorphism (SNP) and how is it used in genomic studies?

B9

How does whole-genome duplication (polyploidy) differ from single-gene duplication in its evolutionary consequences?

B10

What is synteny and what does conserved synteny between species tell us?

Section C · Critical Thinking · Part 2

Analyze and apply concepts; compare your reasoning to the sample answer.

C1

How can retrotransposon reactivation in somatic cells contribute to cancer, and what normally suppresses retrotransposon activity?

C2

A genome-wide association study (GWAS) identifies a SNP in an intergenic region associated with type 2 diabetes. Why might a non-coding SNP affect disease risk?

C3

A deletion on chromosome 15q11-13 causes Prader-Willi syndrome if inherited from the father but Angelman syndrome if inherited from the mother. Explain this using imprinting.

C4

Explain why copy-number variations (CNVs) can have more severe phenotypic effects than point mutations in the same gene.

C5

The Human Genome Project produced a reference genome, but it does not represent the full genomic diversity of humans. What are the limitations of a single reference genome?

Section D · Interactive Questions · Part 2

Enter your answer and click Check for instant feedback.

D1

Approximately what percentage of the human genome is made up of transposable elements? (round number, use %)

D2

How many Hox gene clusters are found in humans? (number)

D3

LINE-1 encodes a reverse transcriptase. What class of transposable element does LINE-1 belong to? (Class I or Class II)

D4

What non-coding RNA is responsible for X-chromosome inactivation in female mammals? (four letters)

D5

In genomic imprinting, IGF2 is expressed only from which parental allele? (maternal or paternal)