1. Membrane Structure and Function
Chapter 7
Membrane Structure and Function

2. Overview: Life at the Edge
The plasma membrane is the boundary that separates the living cell from its surroundings.
The plasma membrane exhibits selective permeability, allowing some substances to cross it more easily than others.
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3. Cellular membranes are fluid mosaics of lipids and proteins
Phospholipids are the most abundant lipid in the plasma membrane
Phospholipids are amphipathic molecules, containing hydrophobic and hydrophilic regions
The fluid mosaic model states that a membrane is a fluid structure with a “mosaic” of various proteins embedded in it
For the Cell Biology Video Structure of the Cell Membrane, go to Animation and Video Files.
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4. Phospholipids: Amphipathic Molecules
WATER
Hydrophilic
head
Figure 7.2 Phospholipid bilayer (cross section)
Hydrophobic
tail
WATER

5. In 1935, Hugh Davson and James Danielli proposed a sandwich model in which the phospholipid bilayer lies between two layers of globular proteins.
Later studies found problems with this model, particularly the placement of membrane proteins, which have hydrophilic and hydrophobic regions.
In 1972, J. Singer and G. Nicolson proposed that the membrane is a mosaic of proteins dispersed within the bilayer, with only the hydrophilic regions exposed to water.
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6. The Fluid Mosaic Model Singer and Nicholson: Phospholipid bilayer
Figure 7.3 The fluid mosaic model for membranes
Hydrophobic regions
of protein
Hydrophilic
regions of protein

7. Freeze-fracture studies of the plasma membrane supported the fluid mosaic model.
Freeze-fracture is a specialized preparation technique that splits a membrane along the middle of the phospholipid bilayer.
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8. Freeze Fracture Method
TECHNIQUE
RESULTS
Extracellular
layer
Proteins
Inside of extracellular layer
Knife
Figure 7.4 Freeze-fracture
Plasma membrane
Cytoplasmic layer
Inside of cytoplasmic layer

9. The Fluidity of Membranes
Phospholipids in the plasma membrane can move within the bilayer.
Most of the lipids, and some proteins, drift laterally.
Rarely does a molecule flip-flop transversely across the membrane/
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10. Fluidity of Membranes Lateral movement (~107 times per second)
Flip-flop
(~ once per month)
(a)Lateral Movement of phospholipids
Fluid
Viscous
Unsaturated hydrocarbon
tails with kinks
Saturated hydro-
carbon tails
(b) Membrane fluidity
Figure 7.5 The
Cholesterol
(c) Cholesterol within the animal cell membrane

11. Hybrid cell Do membrane proteins move? Membrane proteins
RESULTS
Membrane proteins
Mixed proteins
after 1 hour
Mouse cell
Figure 7.6
Human cell
Hybrid cell

12. As temperatures cool, membranes switch from a fluid state to a solid state.
The temperature at which a membrane solidifies depends on the types of lipids.
Membranes rich in unsaturated fatty acids are more fluid that those rich in saturated fatty acids.
Membranes must be fluid to work properly; they are usually about as fluid as salad oil.
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13. Unsaturated hydrocarbon Saturated hydrocarbon tails
Fluidity of Membranes
Fluid
Viscous
Unsaturated hydrocarbon
tails with kinks
Saturated hydrocarbon tails
Figure 7.5b The
(b) Membrane fluidity

14. The steroid cholesterol provides BOTH membrane fluidity and stability.
Cholesterol has different effects on membrane fluidity at different temperatures.
At warm temperatures (such as 37°C), cholesterol restrains movement of phospholipids.
At cool temperatures, it maintains fluidity by preventing tight packing.
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15. Fluidity of Membranes Cholesterol
Figure 7.5c The
(c) Cholesterol within the animal cell membrane

16. Membrane Proteins and Their Functions
A membrane is composed of different proteins embedded in the fluid matrix of the phospholipid bilayer.
Membrane proteins determine most of the membrane’s specific functions.
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17. The detailed structure of an animal cell’s plasma membrane. 7-7
Fibers of
extracellular
matrix (ECM)
Glyco-
protein
Carbohydrate
Glycolipid
EXTRACELLULAR
SIDE OF
MEMBRANE
Figure 7.7
Cholesterol
Microfilaments
of cytoskeleton
Peripheral
proteins
Integral
protein
CYTOPLASMIC SIDE
OF MEMBRANE

18. Peripheral proteins are bound to the surface of the membrane.
Integral proteins penetrate the hydrophobic core.
Integral proteins that span the membrane are called transmembrane proteins.
The hydrophobic regions of an integral protein consist of one or more stretches of nonpolar amino acids, often coiled into alpha helices.
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19. The structure of a transmembrane protein
EXTRACELLULAR
SIDE
N-terminus
Figure 7.8
C-terminus
CYTOPLASMIC
SIDE
 Helix

20. Six major functions of membrane proteins:
Transport
Enzymatic activity
Signal transduction
Cell-cell recognition
Intercellular joining
Attachment to the cytoskeleton and extracellular matrix (ECM)
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21. Membrane Proteins Major Functions:
Signaling molecule
Enzymes
Receptor
ATP
Signal transduction
(a) Transport
(b) Enzyme activity
(c) Signal transduction
Figure 7.9
Glyco-
protein
(e) Intercellular joining
(d) Cell-cell recognition:
Glyco-proteins
( f) Attachment to
the cytoskeleton
and extracellular
matrix (ECM)
Membrane Proteins Functions

22. The Role of Membrane Carbohydrates in Cell-to-Cell Recognition:
Cells recognize each other by binding to surface molecules, often carbohydrates, on the plasma membrane ECM.
Membrane carbohydrates may be covalently bonded to lipids (forming glycolipids) or more commonly to proteins (forming glycoproteins)
Oligosaccharides are short carbohydrate chains on the external side of the plasma membrane.
These vary among species, individuals, and even cell types in an individual.
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23. Synthesis and Sidedness of Membranes
Membranes have distinct inside and outside faces.
The asymmetrical distribution of proteins, lipids, and associated carbohydrates in the plasma membrane is determined when the membrane is built by the ER and Golgi apparatus.
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24. Synthesis of membrane components and their orientation on
the resulting ER 1
Transmembrane
glycoproteins
Secretory
protein
Glycolipid
Golgi
apparatus 2
Vesicle
Figure 7.10 3
Plasma membrane:
Cytoplasmic face 4
Extracellular face
Transmembrane
glycoprotein
Secreted
protein
Membrane glycolipid

25. Membrane structure results in selective permeability
A cell must exchange materials with its surroundings, a process controlled by the plasma membrane.
Plasma membranes are selectively permeable, regulating the cell’s molecular traffic in order to maintain homeostasis.
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26. The Permeability of the Lipid Bilayer
Hydrophobic (nonpolar) molecules, such as hydrocarbons, can dissolve in the lipid bilayer and pass through the membrane rapidly.
Polar molecules, such as sugars, do not cross the membrane easily.
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27. Transport Proteins
Transport proteins allow passage of hydrophilic substances across the membrane.
Some transport proteins, called channel proteins, have a hydrophilic channel that certain molecules or ions can use as a tunnel.
Channel proteins called aquaporins facilitate the passage of water.
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28. A transport protein is specific for the substance it moves.
Other transport proteins, called carrier proteins, bind to molecules and change shape to shuttle them across the membrane.
A transport protein is specific for the substance it moves.
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29. Passive transport is diffusion of a substance across a membrane with no energy investment
Diffusion is the tendency for molecules to spread out evenly into the available space: from High to Low concentrations
Although each molecule moves randomly, diffusion of a population of molecules may exhibit a net movement in one direction until dynamic equilibrium is reached.
At dynamic equilibrium, as many molecules cross one way as cross in the other direction. There is movement, but no net change.
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30. Membrane (cross section)
Diffusion
Molecules of dye
Membrane (cross section)
WATER
Net diffusion
Net diffusion
Equilibrium
(a) Diffusion of one solute
Figure 7.11 The diffusion of solutes across a membrane
Net diffusion
Net diffusion
Equilibrium
Net diffusion
Net diffusion
Equilibrium
(b) Diffusion of two solutes

31. Substances diffuse down their concentration gradient, the difference in concentration of a substance from one area to another
No work must be done to move substances down the concentration gradient
The diffusion of a substance across a biological membrane is passive transport because it requires no cell energy.
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32. Effects of Osmosis on Water Balance
Osmosis is the diffusion of water across a selectively permeable membrane.
Water diffuses across a membrane from the region of lower solute concentration to the region of higher solute concentration.
Water flows down its concentration gradient.
More solute means less (solvent) water in that area.
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33. Less Solute means MORE Water More Solute means LESS water;
Lower
concentration
of solute (sugar)
Higher concentration
of sugar
Same concentration
of sugar
H2O
Selectively
permeable
membrane
Osmosis
Water moves
from H --> L
Figure 7.12 Osmosis
Osmosis

34. Water Balance of Cells Without Walls
Tonicity (solute concentration) is the ability of a solution to cause a cell to gain or lose water.
Isotonic solution: Solute concentration is the same as that inside the cell; no net water movement across the plasma membrane.
Hypertonic solution: Solute concentration is greater than that inside the cell; cell loses water. Cell shrinks.
Hypotonic solution: Solute concentration is less than that inside the cell; cell gains water.
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35. Osmosis: Three Types of Solutions
Hypotonic solution
Isotonic solution
Hypertonic solution
H2O
H2O
H2O
H2O
(a) Animal
cell
Normal
Shriveled
Lysed
H2O
H2O
H2O
H2O
Figure 7.13 The water balance of living cells
(b) Plant
cell
Turgid (normal)
Flaccid
Plasmolyzed

36. Hypertonic or hypotonic environments create osmotic problems for organisms.
Osmoregulation, the control of water balance, is a necessary adaptation for life in such environments.
The protist Paramecium, which is hypertonic to its freshwater environment, has an adaptation: a contractile vacuole that acts as a water pump.
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37. Contractile vacuole of Paramecium: an evolutionary adaptation for osmoregulation
Filling vacuole
50 µm
(a) A contractile vacuole fills with fluid that enters from
a system of canals radiating throughout the cytoplasm.
Contracting vacuole
Figure 7.14
(b) When full, the vacuole and canals contract, expelling
fluid from the cell.

38. Water Balance of Cells with Walls
Cell walls help maintain water balance.
A plant cell in a hypotonic solution swells until the cell wall opposes uptake; the cell experiences turgor (firm).
If a plant cell and its surroundings are isotonic, there is no net movement of water into the cell; the cell becomes flaccid (limp).
In a hypertonic solution, the plant cell looses water, shrivels and experiences plasmolysis.
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39. Facilitated Diffusion: Passive Transport Aided by Proteins
In facilitated diffusion, transport proteins speed the passive movement of molecules across the plasma membrane.
Channel proteins provide corridors that allow a specific molecule or ion to cross the membrane.
Channel proteins include
Aquaporins, for facilitated diffusion of water
Ion channels that open or close in response to a stimulus (gated channels).
For the Cell Biology Video Water Movement through an Aquaporin, go to Animation and Video Files.
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

40. Solute Channel protein Carrier protein
Carrier proteins undergo a subtle change in shape
that translocates the solute-binding site across the membrane
EXTRACELLULAR FLUID
Solute
Channel protein
CYTOPLASM
A channel protein
Figure 7.15 Two types of transport proteins that carry out facilitated diffusion
Solute
Carrier protein
A carrier protein: shape change

41. L --> H pumps solutes.
Active transport uses cell energy to move solutes against their gradients
Facilitated diffusion is still passive because the solute moves down its concentration gradient from a High to Low concentration.
BUT some transport proteins use cell energy to move solutes against their concentration gradients; from a Low to High concentration.
This uphill move is called active transport.
L --> H pumps solutes.
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42. The Need for Energy in Active Transport
Active transport moves substances against their concentration gradient / uphill.
Active transport requires energy, usually in the form of ATP
Active transport is performed by specific membrane carrier proteins that use ATP energy to change shape, thereby pumping the solute across the membrane.
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43. The sodium-potassium pump is one type of active transport system.
Active transport allows cells to maintain concentration gradients that differ from their surroundings.
The sodium-potassium pump is one type of active transport system.
For the Cell Biology Video Na+/K+ATPase Cycle, go to Animation and Video Files.
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44. 1 2 3 6 5 4 EXTRACELLULAR FLUID [Na+] high Na+ [K+] low Na+ Na+ Na+
ATP
[Na+] low P
Na+ P
CYTOPLASM
[K+] high
ADP 1 2 3
Carrier Proteins: Active Transport K+
Figure 7.16, 1–6 The sodium-potassium pump: a specific case of active transport K+ K+ K+ K+ P K+ P 6 5 4

45. Review: Passive transport Active transport Diffusion
ATP
Diffusion
Facilitated diffusion
Figure 7.17 Review: passive and active transport

46. How Ion Pumps Maintain Membrane Potential
Membrane potential is the voltage difference across a membrane.
Voltage is created by differences in the distribution of positive + and negative - ions on the two sides of the membrane.
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47. Two combined forces, collectively called the electrochemical gradient, drive the diffusion of ions across a membrane:
A chemical force (the ion’s concentration gradient)
An electrical force (the effect of the membrane potential on the ion’s movement)
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48. An electrogenic pump is a transport protein that generates voltage across a membrane.
The sodium-potassium pump (Na+K+ pump) is the major electrogenic pump of animal cells.
A key electrogenic pump driving chemiosmosis in many cells is a proton pump.
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49. An Electrogenic Pump Proton pump – EXTRACELLULAR FLUID + ATP – + H+ H+
Figure 7.18 An – +
CYTOPLASM H+ – +

50. Cotransport: Coupled Transport by a Membrane Protein
Cotransport occurs when active transport of a solute indirectly drives the transport of another solute.
Plants commonly use the gradient of hydrogen ions generated by proton pumps to drive active transport of nutrients into the cell.
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51. – + H+ ATP H+ – + H+ H+ – + H+ H+ – + H+ H+ – + – + Diffusion of H+
Cotransport: active transport driven by a concentration gradient – + H+
ATP H+ – + H+
Proton pump H+ – + H+ H+ – + H+
Diffusion
of H+
Sucrose-H+
cotransporter
Figure 7.19 H+ –
Sucrose + – +
Sucrose

52. Bulk transport across the plasma membrane occurs by exocytosis and endocytosis
Small molecules and water enter or leave the cell through the lipid bilayer or by transport proteins.
Large molecules, such as polysaccharides and proteins, cross the membrane in bulk via vesicles.
Bulk transport requires energy.
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53. Exocytosis: LARGE molecules OUT
In exocytosis, transport vesicles migrate to the membrane, fuse with it, and release their contents outside the cell.
Many secretory cells use exocytosis to export their products.
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54. Endocytosis: LARGE molecules IN
In endocytosis, the cell takes in macromolecules by forming vesicles from the plasma membrane.
Endocytosis is a reversal of exocytosis, involving different proteins.
There are three types of endocytosis:
Phagocytosis (“cellular eating”)
Pinocytosis (“cellular drinking”)
Receptor-mediated endocytosis
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55. The vacuole then fuses with a lysosome to digest the particle.
In phagocytosis a cell engulfs a particle as the plasma membrane projects pseudopods which surround the particle, forming a vacuole.
The vacuole then fuses with a lysosome to digest the particle.
For the Cell Biology Video Phagocytosis in Action, go to Animation and Video Files.
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56. Endocytosis in animal cells PHAGOCYTOSIS PINOCYTOSIS
EXTRACELLULAR
FLUID
CYTOPLASM
1 µm
Pseudopodium
Pseudopodium
of amoeba
“Food”or
other particle
Bacterium
Food
vacuole
Food vacuole
An amoeba engulfing a bacterium
via phagocytosis (TEM)
PINOCYTOSIS
0.5 µm
Plasma
membrane
Pinocytosis vesicles
forming (arrows) in
a cell lining a small
blood vessel (TEM)
Vesicle
RECEPTOR-MEDIATED ENDOCYTOSIS
Coat protein
Receptor
Coated
vesicle
Figure 7.20
Coated
pit
Ligand
A coated pit
and a coated
vesicle formed
during
receptor-
mediated
endocytosis
(TEMs)
Coat
protein
Plasma
membrane
0.25 µm

57. Endocytosis in animal cells—phagocytosis
EXTRACELLULAR
FLUID
CYTOPLASM
1 µm
Pseudopod
Pseudopodium
of amoeba
“Food” or
other particle
Bacterium
Food
vacuole
Figure 7.20
Food vacuole
An amoeba engulfing a bacterium
via phagocytosis (TEM)

58. Pinocytosis is referred to as “cell drinking.”
In pinocytosis, molecules are taken in when extracellular fluid is “gulped,” surrounded by plasma membrane forming tiny vesicles.
Pinocytosis is referred to as “cell drinking.”
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59. Endocytosis in animal cells—pinocytosis
Plasma
membrane
Pinocytosis vesicles
forming (arrows) in
a cell lining a small
blood vessel (TEM)
Vesicle
Figure 7.20

60. In receptor-mediated endocytosis, binding of ligands to receptors triggers vesicle formation.
A ligand is any molecule that binds specifically to a receptor site of another molecule using shape match.
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61. Receptor-Mediated Endocytosis
Coat protein
Receptor
Coated
vesicle
Coated
pit
Ligand
A coated pit
and a coated
vesicle formed
during
receptor-
mediated
endocytosis
(TEMs)
Coat
protein
Figure 7.20 Endocytosis in animal cells—receptor-mediated endocytosis
Plasma
membrane
0.25 µm

62. Review: Passive transport: Facilitated diffusion Channel Carrier
protein
Carrier
protein

63. Review:
Active transport:
ATP

64. Review: Diffusion Osmosis
Environment:
0.01 M sucrose
0.01 M glucose
0.01 M fructose
“Cell”
0.03 M sucrose
0.02 M glucose

65. Diffusion of Multiple Molecules

66. You should now be able to:
Define the following terms: amphipathic molecules, aquaporins, diffusion.
Explain how membrane fluidity is influenced by temperature and membrane composition.
Distinguish between the following pairs or sets of terms: peripheral and integral membrane proteins; channel and carrier proteins; osmosis, facilitated diffusion, and active transport; hypertonic, hypotonic, and isotonic solutions.
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67. Explain how transport proteins facilitate diffusion.
Explain how an electrogenic pump creates voltage across a membrane, and name two electrogenic pumps.
Explain how large molecules are transported across a cell membrane.
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