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Chapter 10

Membrane Structure

Molecular Biology of the Cell 7e · Part 1 of 2 · 10 MCQs per part · 60 total questions
Part 1 of 2
The Lipid Bilayer and Membrane Composition
Every cell is bounded by a membrane that both defines it and controls what enters and leaves. Understanding how a simple sheet of lipid molecules, just 5 nm thick, can carry out the extraordinary range of functions attributed to biological membranes is one of the triumphs of twentieth-century cell biology.

10.1 The Lipid Bilayer

Biological membranes are built from phospholipid bilayers. Phospholipids are amphipathic molecules with a hydrophilic glycerophosphate head and two hydrophobic fatty acid tails. In aqueous solution they spontaneously self-assemble into bilayers, with tails sequestered from water and heads exposed.

The three major membrane lipid classes are: glycerophospholipids (e.g., phosphatidylcholine, phosphatidylserine), sphingolipids (e.g., sphingomyelin), and sterols (cholesterol in animal cells).

Key term
Amphipathic molecule

A molecule with both hydrophilic (water-loving) and hydrophobic (water-fearing) regions; the amphipathic nature of phospholipids drives bilayer self-assembly.

Bilayer thickness is approximately 5 nm. The bilayer is impermeable to most ions and polar molecules but freely permeable to small nonpolar molecules (O₂, CO₂, N₂) and small uncharged polar molecules (H₂O, urea).

10.2 Membrane Fluidity and Cholesterol

Membrane fluidity is determined by lipid composition. Unsaturated fatty acids (cis double bonds introduce kinks) increase fluidity; longer saturated chains decrease fluidity. Cholesterol has a dual role: it broadens the temperature range over which the bilayer is fluid, preventing both crystallisation at low temperature and excess fluidity at high temperature.

The transition temperature (Tm) separates the ordered gel phase from the disordered fluid (liquid-crystalline) phase.

Pause & Recall
What is the role of cholesterol in regulating membrane fluidity?
Cholesterol intercalates between phospholipids, restraining movement at high temperatures (reducing fluidity) and preventing tight packing at low temperatures (preventing gel formation), so it buffers fluidity across physiological temperature ranges.

10.3 The Fluid Mosaic Model

The Singer-Nicolson fluid mosaic model (1972) describes proteins as embedded in or associated with a fluid lipid bilayer, free to diffuse laterally unless constrained.

FRAP (fluorescence recovery after photobleaching) directly demonstrates lateral diffusion of lipids and some membrane proteins by bleaching a patch of fluorescently labelled membrane and monitoring recovery as unbleached molecules diffuse in.

Lipid flip-flop (transverse diffusion) is extremely slow (t½ ~ days) for phospholipids in the absence of enzymes. Flippases (ATP-driven, move PS/PE to inner leaflet), floppases (move PC/SM to outer leaflet), and scramblases (bidirectional, Ca²⁷-activated) maintain lipid asymmetry.

10.4 Membrane Asymmetry

The two leaflets of the plasma membrane have distinct lipid compositions. In the outer leaflet: phosphatidylcholine (PC), sphingomyelin (SM), glycolipids. In the inner leaflet: phosphatidylserine (PS), phosphatidylethanolamine (PE), PI(4,5)P₂.

PS exposure on the outer leaflet is a signal for apoptosis (recognition by macrophages) and platelet activation.

Key term
Flippase

A P4-type ATPase that uses ATP hydrolysis to flip specific phospholipids (PS, PE) from the outer to the inner leaflet, establishing and maintaining bilayer asymmetry.

Practice Questions — Part 1Score: 0 / 10

1. Which of the following best describes the structure of a phospholipid?

2. The transition temperature (Tm) of a lipid bilayer would be LOWEST for membranes containing:

3. What experimental technique directly demonstrated the lateral diffusion of membrane proteins?

4. Which lipid is found predominantly in the INNER leaflet of the plasma membrane?

5. Cholesterol's effect on membrane fluidity can best be described as:

6. Which enzyme class moves phospholipids from the outer to the inner leaflet of the plasma membrane using ATP?

7. Which of the following can cross a pure phospholipid bilayer most rapidly by simple diffusion?

8. The fluid mosaic model proposed by Singer and Nicolson describes biological membranes as:

9. Exposure of phosphatidylserine on the outer leaflet of a cell's plasma membrane is a signal for:

10. Which of the following is NOT a property of glycerophospholipids?

Part 1 complete! Score: 0 / 10

Section B · Recall Questions · Part 1

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

B1

What forces drive phospholipid self-assembly into a bilayer?

B2

Describe how unsaturated fatty acids affect membrane fluidity.

B3

What is FRAP and what does it measure?

B4

Explain membrane lipid asymmetry — which lipids are in each leaflet?

B5

What distinguishes a flippase from a scramblase?

B6

Why are ions such as Na⁺ unable to cross a lipid bilayer by simple diffusion?

B7

What is the role of cholesterol in the plasma membrane of animal cells?

B8

Describe the structure of sphingomyelin and its enrichment in lipid rafts.

B9

What structural feature of cis-unsaturated fatty acids makes membranes more fluid?

B10

How does the lipid composition of the inner mitochondrial membrane differ from the plasma membrane, and why?

Section C · Critical Thinking · Part 1

Develop analytical responses, then compare with the sample.

C1

A researcher discovers a new antibiotic that inserts into bacterial phospholipid bilayers and creates non-specific pores. Explain why this mechanism might selectively kill bacteria without harming human cells.

C2

FRAP experiments on a membrane protein show 40% fluorescence recovery after 10 minutes. Interpret these results in terms of membrane protein mobility.

C3

How might increasing dietary intake of omega-3 polyunsaturated fatty acids affect the plasma membranes of neurons, and why might this be beneficial?

C4

Explain why PS exposure on the outer leaflet is a reliable signal for apoptosis, given that PS is normally confined to the inner leaflet.

C5

Design an experiment to test whether a newly identified protein is an integral or peripheral membrane protein.

Section D · Interactive Questions · Part 1

Enter your answer and click Check for instant feedback.

D1

The process by which lipids spontaneously move from the outer to the inner leaflet using ATP is called ________.

D2

Small nonpolar molecules like O₂ cross the lipid bilayer by ___________ diffusion.

D3

The experimental model describing proteins embedded in a fluid lipid bilayer is the ________ model.

D4

The lipid that both prevents gel formation at low temperatures and restricts excess fluidity at high temperatures is ___________.

D5

Bilayer lipid asymmetry is broken down during apoptosis by enzymes called ___________.
Part 2 →

Having established the lipid foundation of biological membranes, we now turn to the proteins that give membranes their functional diversity — from transmembrane channels to GPI-anchored receptors — and examine how cells organise membrane components into specialised domains.

Part 2 of 2
Membrane Proteins, Dynamics, and Specialised Domains

10.5 Types of Membrane Proteins

Membrane proteins are classified as integral (transmembrane or monotopic) or peripheral.

Integral transmembrane proteins span the bilayer via hydrophobic α-helices (most common) or β-barrel structures (found in outer membranes of gram-negative bacteria and mitochondria/chloroplasts).

Glycosylphosphatidylinositol (GPI)-anchored proteins are attached to the outer leaflet via a lipid anchor; they lack a cytoplasmic domain and are enriched in lipid rafts.

Peripheral membrane proteins associate non-covalently with the membrane surface (via electrostatic or hydrophobic interactions) or with transmembrane proteins.

Key term
Transmembrane domain

A stretch of ~20 hydrophobic amino acids that forms an α-helix and spans the bilayer; integral membrane proteins typically contain one or more transmembrane helices.

10.6 Lipid Rafts

Lipid rafts are small (10–200 nm), dynamic, ordered microdomains in the outer leaflet enriched in cholesterol, sphingomyelin, and GPI-anchored proteins. They concentrate signalling molecules and receptors, potentially facilitating signal transduction.

Pause & Recall
What is the evidence for lipid rafts?
Biochemical: some membrane proteins fractionate into detergent-resistant membranes (DRMs) with cholesterol and sphingomyelin. Imaging: super-resolution microscopy reveals nanoscale protein clusters. Functional: depletion of cholesterol (with methyl-β-cyclodextrin) disrupts signalling. However, DRMs may be artefacts of detergent treatment, and in-vivo raft size/lifetime is debated.

10.7 Protein Mobility and Membrane Domains

Not all membrane proteins diffuse freely; many are corralled into domains by interactions with the underlying cortical cytoskeleton (membrane skeleton). The 'picket-fence' model proposes that transmembrane proteins anchored to the cytoskeleton act as pickets, and actin-filament corrals act as fences, restricting lateral diffusion of membrane proteins and lipids.

Cell-cell junctions (tight junctions, gap junctions) further restrict membrane protein mobility and create distinct apical and basolateral membrane domains in epithelial cells.

10.8 Membrane Permeability and Biological Functions

The selective permeability of biological membranes underpins cellular homeostasis. While the bilayer itself bars passage of ions, sugars, and amino acids, specialised membrane proteins (channels, carriers, pumps) provide pathways.

Membrane curvature is actively generated and sensed: BAR domain proteins sense or induce curvature; this is important for vesicle budding and tubulation.

Membrane composition is tightly regulated; defects in membrane lipid metabolism cause diseases (e.g., Niemann-Pick disease from sphingomyelinase deficiency; Tangier disease from ABCA1 cholesterol transporter defects).

Practice Questions — Part 2Score: 0 / 10

1. Which structural feature characterises most integral membrane proteins?

2. GPI-anchored proteins are exclusively found on which leaflet of the plasma membrane?

3. Which of the following best describes the 'picket-fence' model of membrane organisation?

4. Depletion of cholesterol from the plasma membrane using methyl-β-cyclodextrin would be expected to:

5. How do epithelial cells maintain distinct apical and basolateral membrane domains?

6. A β-barrel protein structure in a biological membrane is most likely found in:

7. A peripheral membrane protein can be selectively removed from membrane preparations by:

8. Niemann-Pick disease type A/B is caused by a deficiency of which enzyme?

9. Which domain of peripheral membrane proteins is responsible for sensing membrane curvature?

10. Which disease is caused by defects in the ABCA1 transporter that exports cholesterol from cells?

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 two structural differences between integral and peripheral membrane proteins.

B2

What is a GPI anchor and how is it added to a protein?

B3

What are lipid rafts and what proteins do they concentrate?

B4

Explain how tight junctions maintain membrane polarity in epithelial cells.

B5

What is the 'picket-fence' model of membrane protein diffusion?

B6

Describe the main evidence used to identify lipid rafts.

B7

How do BAR domain proteins generate membrane curvature?

B8

What is the clinical significance of phosphatidylserine exposure in platelets?

B9

Describe the structural basis for the β-barrel architecture of outer-membrane proteins compared with those in mitochondria.

B10

What disease results from deficiency of acid sphingomyelinase, and what accumulates?

Section C · Critical Thinking · Part 2

Develop analytical responses, then compare with the sample.

C1

A mutant cell line lacks functional flippases. Predict the consequences for membrane lipid asymmetry and cell biology.

C2

Why might depletion of cholesterol from the membrane affect the function of a receptor tyrosine kinase located in a lipid raft?

C3

Explain how a GPI-anchored protein on the outer leaflet could influence signalling on the inner leaflet, given that it has no transmembrane or cytoplasmic domain.

C4

In epithelial cells of the kidney tubule, Na⁺/K⁺-ATPase pumps are restricted to the basolateral membrane. What structural mechanisms ensure this polarised distribution?

C5

How might a mutation that converts a charged residue in the interior of a lipid raft to a hydrophobic one affect raft association of that protein?

Section D · Interactive Questions · Part 2

Enter your answer and click Check for instant feedback.

D1

Proteins anchored to the outer leaflet via a phosphoinositol-glycan linkage are called ___________ proteins.

D2

The crescent-shaped protein domain that senses and induces membrane curvature is the ___________ domain.

D3

In gram-negative bacteria, outer membrane channels formed by β-barrel proteins that allow passive diffusion of small molecules are called ___________.

D4

The disease caused by deficiency of acid sphingomyelinase is called ___________ disease.

D5

The junction type that forms a diffusion fence between apical and basolateral membrane domains in epithelia is the ___________ junction.