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Biochemistry Unit 7: Membranes and Transport

Membranes separate the cell from its surroundings and control what enters and leaves. This unit explains bilayer structure, membrane proteins, the kinds of transport, and how to calculate the energy of moving solutes and ions.

Covers: Fluid mosaic model and bilayer properties, Membrane proteins, Simple and facilitated diffusion, Channels and selectivity, Primary active transport, Secondary active transport, Electrochemical gradients and membrane potential

Study guide

Biological membranes follow the fluid mosaic model: a lipid bilayer, about 5 nm thick, in which proteins float. Lipids diffuse quickly sideways within a leaflet, as FRAP experiments show, but rarely flip between leaflets without flippase enzymes, so the two leaflets differ in composition. Fluidity depends on lipid tails: cis-unsaturated and shorter chains increase fluidity, while saturated and longer chains decrease it. Cholesterol buffers fluidity, stiffening warm membranes and preventing cold ones from gelling. Small nonpolar molecules such as O₂ and CO₂ diffuse straight through; ions and large polar molecules need proteins.

Membrane proteins are integral or peripheral. Integral proteins cross the bilayer, often with α-helices of about 20 hydrophobic residues, or as β-barrels, and need detergents to extract. Peripheral proteins bind the surface through ionic and polar contacts and are released by salt or pH changes. Lipid-anchored proteins are tethered by covalent fatty acyl, prenyl or GPI anchors. Transport proteins come in two broad types: channels, which form pores that let ions or water flow quickly down their gradients, and transporters (carriers), which bind solutes and change shape by alternating access, showing saturation kinetics.

Passive transport moves solutes down their electrochemical gradient. Simple diffusion requires no protein and rises linearly with concentration. Facilitated diffusion uses a channel or carrier, such as GLUT1 for glucose or aquaporins for water. Ion channels can be highly selective: the K⁺ channel's selectivity filter uses backbone carbonyl oxygens that mimic K⁺'s water shell, so K⁺ passes about 1,000 times more easily than the smaller Na⁺. Many channels are gated by voltage, ligands or mechanical force. Carriers such as GLUT1 behave like enzymes: their rate follows v = Vmax[S]/(Kt + [S]), where Kt is the solute level giving half-maximal rate, and they can be competitively inhibited by similar molecules.

Active transport moves solutes uphill. Primary active transport uses ATP directly: P-type ATPases such as the Na⁺/K⁺-ATPase (3 Na⁺ out, 2 K⁺ in per ATP) and Ca²⁺-ATPases, plus ABC transporters. Because it moves net positive charge out, the Na⁺/K⁺ pump is electrogenic. Secondary active transport uses an ion gradient made by a primary pump. SGLT1 symports 2 Na⁺ with each glucose into intestinal cells, and the Na⁺/Ca²⁺ exchanger antiports Na⁺ in and Ca²⁺ out. Drugs such as ouabain and digoxin inhibit the Na⁺/K⁺ pump, raising cell Na⁺ and, in the heart, Ca²⁺.

The free energy of transport is ΔG = RT ln(C_in/C_out) + zFΔψ for moving a solute into the cell, where Δψ is the inside potential relative to outside. For uncharged solutes only the concentration term applies; a tenfold gradient is worth about 5.9 kJ/mol at 37 °C. For ions, the membrane potential matters: with the inside at −70 mV, cations are pulled in. The Nernst equation, E = (RT/zF) ln([out]/[in]), gives the potential at which an ion is at equilibrium. The resting potential lies near the K⁺ Nernst potential because resting membranes are most permeable to K⁺.

Key ideas

Formulas and rules

ΔG (moving in) = RT ln(C_in/C_out) + zFΔψ, with Δψ = ψ_in − ψ_out
RT ≈ 2.58 kJ/mol at 37 °C; F = 96.485 kJ/(V·mol)
Nernst: E = (RT/zF) ln([out]/[in]) ≈ (26.7 mV/z) ln([out]/[in]) at 37 °C
Carrier kinetics: v = Vmax[S]/(Kt + [S])
Na⁺/K⁺-ATPase: 3 Na⁺ out + 2 K⁺ in per ATP
Max accumulation by coupling: RT ln(ratio) = energy supplied by the driving ions

Common mistakes

Before the test, make sure you can

Words to know

Fluid mosaic model
A model of membranes as fluid lipid bilayers with mobile embedded proteins.
Flippase
An enzyme that moves lipids between membrane leaflets to keep asymmetry.
Integral protein
A protein embedded in or spanning the bilayer, removable only with detergent.
Peripheral protein
A protein bound loosely to the membrane surface, removable by salt or pH changes.
Channel
A pore protein allowing rapid flow of specific ions or water down their gradient.
Transporter (carrier)
A protein that binds a solute and changes shape to move it, showing saturation.
Facilitated diffusion
Protein-mediated movement down a gradient with no energy input, such as GLUT1.
Primary active transport
Uphill transport powered directly by ATP hydrolysis.
Secondary active transport
Uphill transport powered by an ion gradient, such as SGLT1.
Symport
Movement of two solutes in the same direction by one transporter.
Antiport
Movement of two solutes in opposite directions by one transporter.
Na⁺/K⁺-ATPase
A P-type pump exporting 3 Na⁺ and importing 2 K⁺ per ATP.
Electrogenic
Producing net charge movement across a membrane.
Nernst potential
The membrane voltage at which a given ion is at equilibrium.
Resting membrane potential
The steady voltage across a resting cell membrane, about −70 mV in neurons.
Selectivity filter
The narrow part of a channel that determines which ion can pass.
Aquaporin
A channel protein that conducts water rapidly while excluding protons.
FRAP
Fluorescence recovery after photobleaching, used to measure lateral diffusion.

Sample practice questions

30 more questions like these, from level 1 to 5, are in the practice set.

Question 1 (level 1)

The fluid mosaic model describes membranes as:

  1. A lipid bilayer with proteins that can move laterally
  2. A single layer of phospholipids with heads inward
  3. A rigid protein layer coated with lipids
  4. A lipid bilayer sandwiched between two protein sheets
Show the answer and steps

Answer: A) A lipid bilayer with proteins that can move laterally

  1. Singer and Nicolson (1972) proposed a fluid bilayer with embedded and attached proteins.
  2. Lipids and many proteins diffuse sideways within each leaflet.
  3. The protein-sandwich idea was the older Davson-Danielli model.
Question 2 (level 1)

Which molecule crosses a pure lipid bilayer most easily by simple diffusion?

  1. ATP
  2. Na⁺
  3. O₂
  4. Glucose
Show the answer and steps

Answer: C) O₂

  1. Small, nonpolar molecules dissolve in the hydrocarbon core and pass through.
  2. O₂ and CO₂ cross quickly.
  3. Ions, sugars and charged nucleotides need transport proteins.
Question 3 (level 3)

At 37 °C with [K⁺] = 140 mM inside and 5 mM outside, what is the K⁺ equilibrium (Nernst) potential? (RT/F = 26.7 mV)

  1. −89 mV
  2. −67 mV
  3. +89 mV
  4. −39 mV
Show the answer and steps

Answer: A) −89 mV

  1. E = (RT/zF) ln([out]/[in]) = 26.7 mV × ln(5/140).
  2. ln(0.0357) = −3.33, so E = −89 mV.
  3. +89 inverts the ratio; −39 uses log₁₀ without the 2.303 factor.